Wildlife Responses to Pedestrians and Dogs

Scott G Miller Richard L Knight Clinton K Miller

Wildlife Society Bulletin Vol 29 No 1 (Spring 2001) pp 124-132

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PEDESTRIANS AND DOGS

Wildlife responses to pedestrians and dogs

Scott G llilZer Rich clrd L Knight and Clinton K lliZler

Abstract As participation in outdoor recreational activities escalates land managers struggle to develop management policies that ensure coexistence of wildlife and recreation How-ever this requires an understanding of how wildlife responds to various forms of recre- ational activities and the spatial context in which the activities occur Therefore we measured responses of 2 species of grassland songbirds one species of forest songbird and mule deer (Odocoileus hemionus) exposed to a pedestrian a pedestrian accompa- nied by a dog on leash and a dog alone (only for grassland birds) on and away from recreational trails We assessed the area ot influence for each treatment by determining the probability that an animal would flush or become alert (for mule deer only) given its perpendicular distance to a trail or a line of movement in areas without trails When ani- mals were disturbed we measured flush distance (the distance between the disturbance and the animal when flushed) distance moved and for mule deer alert distance (the dis- tance between the disturbance and the deer when it became alert) For all species area of influence flush distance distance moved and alert distance (for mule deer) was greater when activities occurred off-trail versus on-trail Generally among on-trail and off-trail treatments in grasslands for vesper sparrows (Pooecetes gramineus) and western meadowlarks (Sturnella neglects) the smallest area of influence and shortest flush dis- tance and distance moved resulted from the dog-alone treatment and these responses were greater for the pedestrian-alone and dog-on-leash treatments In forests for Amer- ican robins (Turdus migratorius) the area of influence flush distance and distance moved did not generally differ between the pedestrian-alone and dog-on-leash treatments For mule deer presence of a dog resulted in a greater area of influence alert and flush dis- tance and distance moved than when a pedestrian was alone Natural lands managers can implement spatial and behavioral restrictions in visitor management to reduce dis- turbance by recreational activities on wildlife Restrictions on types of activities allowed in svme areas such as prohibiting dogs or restricting use to trails wi l l aid in minimizing disturbance Additionally managers can restrict the number and spatial arrangement of trails so that sensitive areas or habitats are avoided

Key words American robin disturbance dog mule deer outdoor recreation pedestrian trail vesper sparrow western meadowlark

As participation inoutdoor recreational activities Because outdoor recreation has become common escalates land managers are becoming concerned and widespread managers must n o w incorporate about the effects of recreation o n wildlife (Boyle actions into their management decisions that mini- and Samson 1985 Knight and Gutzwiller 1995) mize potential impacts o f these activities This

Address for Scott C Miller and Richard L Knight Department of Fishery and Wildl i fe B ~ o l o g ~ Colorado State University Fort Collins C O 80523 USA e-mail tor Mil ler scott-g_millerfw~sgovpresent address ior Mil ler United States Fish and Wildl i fe Ser- vice AlamosaIMonte Vista National Wildl i te Refuge 9383 El Rancho Lane Alamosa C O 81 101 USA Address for Clinton K Mil ler Department o i Open Space 66 S Cherryvale Road Boulder C O 80303 USA present address The Nature Conservancy Northern Tallgrass Prairie Ecoregion PO Box 81 6 Clear Lake SD 57226 USA

Wildlife Society Bulletin 2001 29(1)124-132 Peer refereed

Pedestrians and dogs Miller et al 125

requires an understanding of how wildlife responds to various forms of outdoor recreation and also the temporal and spatial context in which the activity occurs (Knight and Cole 1995)

Information on how wildlife reacts to hikers and dogs is limited although preliminary evidence sug-gests that presence of dogs increases the response For example mountain sheep (Ouis ccuzadensis MacArthur et al 19791982)golden plovers (Pluzli-nlis upricaria Yalden and Yalden 1990) and mar-mots (LMarmotcz marmotu Mainini et al 1993) exhibited a greater response when pedestrians were accompanied by a dog compared to solitary pedestrians

Location and frequency of recreational activities also can influence wildlife responses (Knight and Cole 1995) If animals perceive an activity as spa-tially predictable and nonthreatening they may habituate to that activity (Whittaker and Knight 1998) For example humans approaching from a parking area (an area with consistent hunlan use) elicited less of a response from mountain sheep than did humans approaching from over a ridge where human use was sporadic (MacArthur et al 1982)

Of the numerous studies on effects of recreation-al activities on wildlife most present information on flush distance (the distance between the activi-ty and the animal when it flushes) as the animal is approached directly by humans Although bird-watchers photographers and others do approach wildlife most recreationists do not go out of their way to do so Rather most recreationists such as hikers walking on trailsdo not commonly leave the trail To investigate this type of disturbancewe cor-related an animals flush response with its perpen-dicular distance to the trail or line of human move-ment With this informationwe were able to assess an area of influence for each treatment Area of influence was defined as the probability that an animal will flush or become alert (for mule deer only) at a given perpendicular distance from a trail or line of human movement The greater the area of influence the more disturbing the activity is to wildlife For example if the probability of flushing for a bird 30 m away from a trail is 040 to a pedes-trian accompanied by a dog and 070 to a pedestri-an alone then the area of influence is greater for the pedestrian alone

Our objective was to assess the area of influence around a lone pedestrian a pedestrian accompa-nied by a dog on leash and a dog aloneon and off trails For animals that flushedwe compared infor-

mation on flush distance (the distance between the activity and the animal when flushed) and distance moved to further assess the magnitude of distur-bance for each treatment Additionally for mule deer we compared illformation on alert distance (the distance between the activity and the deer when it became alert) among treatments In grass-lands we recorded responses of vesper sparrows and w-estern meadowlarks to all treatments In forests we recorded responses of American robins and mule deer to all treatments except the dog alone For each specieswe tested the null hypoth-esis that the area of influence and magnitude of dls-turbance did not differ between treatments

Methods and study area We conducted our study on 8000 ha of City of

Boulder Open Space property in and around the city of Boulder Colorado (40deg00N 105deg1845E) Elevation within the study area ranged from 1219 to 2438 m encompassing forest riparian shrub-land and grassland habitats Visitor use on City of Boulder Open Space is approximately 2 million vis-itsyear and is greatest during the spring followed by summer fall and winter (Zeller et al 1993) Recreational activities included hiking wildlife viewing exercising pets jogging mountain biking and horseback riding (hunting is not allowed)

We located study sites in pine forests and mixed-grass prairies Forests were dominated by pon-derosa pine (Pinus potzderosa) associated with shrubsgrasses and forbs Mixed-grass prairies con-tained a variety of tall mid-height and shortgrass species including little bluestem (Scbiznch-yriurn scopariurn) western wheatgrass (Agropj~ron smithiz3blue grama (Boz~telouagrcacilis) and side oats grama (Bouteloua curtipendz~la)

We conducted treatments on trails and for off-trail sites on areas gt400 m from trails Trails received frequent use whereas off-trail sites were used sporadically by recreationists We located all sites gt800 nl from urban developmentand gt400 m from physiographic features such as forest edge riparian areas and ridge lines Trail width was 125 k022 m (mean 1 SE) in the grasslands and 11-+ 020 m (mean 1 SE) in the forests

We collected data between 14 April and 20 July 1996 We rotated visits to on-trailand off-trailsites to avoid repeatedly sampling the same areas Birds and mule deer were not marked so we could not assure the same individuals were not mdtiply sampled

126 WWlfe Society Bulletin 200129(1) 124- 132

Grassland In grasslands we recorded responses of vesper

sparrows and western meadowlarks to 3 activities on- and off-trail 1) a pedestrian alone 2) a pedestri- an accompanied by a dog on leash and 3) a dog alone We selected these species because of their abundance on the study site and we were able to obtain adequate sample sizes for statistical compar- isons For treatments involving dogs we used either a 25-kg or a 40-kg dog Leash length was 18 m For on- and off-trail dog alone treatments the dog maintained an approximate distance of 20 m in front of the observer For dog-alone treatments we assumed that birds were responding to the dog only and not the observer In no case did the dogs attempt to chase birds

For on-trail treatments we detected individual birds on or near the trail ahead of us and proceed- ed along the center of the trail at approximately 15 msecond until the bird flushed or the observ- er had passed by eliciting no flush response At that time the observer stopped momentarily to record 1) flush response 2) the perpendicular distance between the bird and the trail 3) flush distance and 4) distance moved On off-trail sites we locat- ed birds on or near our line of movement and pro- ceeded parallel to the birds position so as to pass by at various distances (0 m to 200 m perpendicu- lar distance) After the bird flushed or the observer passed by eliciting no flush response we stopped momentarily to record the same information as that for on-trail treatments

Forest In forests we recorded responses of American

robins and mule deer both on- and off-trail to a pedestrian alone and a pedestrian accompanied by a dog on leash We selected these species because of their abundance on the study site and we were able to obtain adequate sample sizes for statistical comparisons Information for a dog alone was not recorded because we were unable to maintain an adequate distance behind the dog and still assume that robins or deer were responding only to the dog

We conducted treatments with robins and deer the same as in the grassland trials For robins we also measured (to the nearest 1 m) height above the ground (if perched in a tree) For deer we also recorded 1) alert response (ie lifted its head) 2) alert distance and 3) time elapsed from when a deer first exhibited a response until it resumed the

pre-disturbance behavior When group size was gt 1 we recorded information for the first deer to elicit a response We used a Lietz rangefinder (model 3390) to measure all distances to the nearest 1 m

Statistical analyses We used logistic regression (GENMOD proce-

dure SAS Institute Inc 1993) to determine whether flush response (and alert response for deer) of indi- vidual species was correlated with treatment per- pendicular distance to trail or line of movement date time of ampa) height of bird if perched in tree (for American robins) and group size and sex (for mule deer) For the animals that flushed we used analysis of variance (SAS Institute Inc 1988) to compare flush distance among treatments and also distance moved among treatments of individual species Because we attempted to simulate typical recreationist behavior (ie continuing to proceed along the trail or line of movement without stop- ping) many deer remained alert to our presence until we moved out of their sight Consequently mean and SE of time elapsed from when a deer first exhibited a response until it resumed the pre-disturbance activity could not be determined and we did not conduct statistical analysis comparing treatments For each grassland treatment we com- pared flush distance and also distance moved between vesper sparrows and western mead-owlarks using t-tests (SAS Institute Inc 1988) We used an a=005 for all analyses

Results Grassland

We conducted 462 and 393 trials for vesper spar- rows and western meadowlarks respectively For both species logistic regression models indicated that treatment (Plt0001) and perpendicular dis- tance of the bird (Plt0001) to a trail or line of movement (for off-trail) were significant predictors of flush response (Figure 1) The shorter the per- pendicular distance of a bird to the trail or line of movement the greater the probability that a bird would flush For both species the area of influence was greater for off-trail treatments than for on-trail treatments (Figure 1) For vesper sparrows on- and off-trail and also for western meadowlarks on-trail the dog-alone treatment resulted in a smaller area of influence than the pedestrian-alone or dog-on- leash treatments which did not differ from each other For western meadowlarks area of influence

Pedestrians and dogs Miller et al 127

(a) Pedestrian alone (on-trail) Dog on leash (on-trail) Dog alone (on-trail)

Pedestrian alone (off-trail) Dog on leash (off-trail) Dog alone (off-trail)

Pedestrian alone (on-trail) Dog on leash (on-trail) Dog alone (on-trail)(b)

rC

0 5 Pedestrian alone (off-trail) Dog on leash (off-trail)- Dog alone (off-trail)

~ ~~~~ ~ ~~

0 00

Meters from trail (for on-trail) or line of movement (for off-trail) Figure 1 Predicted probability of a vesper sparrow (a) and western meadowlark ibi flushing to treatments in grasslands during 1996 City of Boulder Open Space Boulder Colorado (dashed lines indicate 95 CIj

did not differ among off-trail treatments Date and time of day were not significant predictors of whether a bird would flush (Pgt005 for both species)

When vesper sparrows flushed mean flush dis-tance differed among treatments (F3269= 1175Plt 0001Table 1) Flush distance was greater for the off-trail pedestrian-alone and off-trail dog-on-leash treatments than for any other treatment Other

treatments did not differ For vesper sparrows dis-tance moved did not differ among treatments (Fj2= 146P=0204Table 1) however birds at off-trail sites tended to fly farther when compared to on-trail sites

When western meadowlarks flushed mean flush distance differed among treatments (Fj244=800 Plt0001Table 1) For each activityflush distance was greater for off-trail than on-trail treatments

128 Wildlife Sociep Bulletin 200129(1)124- 132

Table 1 Mean (SEI of flush distancea and distance moved for (t2298 Plt0005) and meadowlarks flew greater vesper sparrows and western meadowlarks in grasslands City distances once flushed (t340~10001) of Boulder Open Space Boulder Colorado 1996

Flush Distance FOfamp Treatment distance (m) moved ~ m We ran 228 trials for American robins Logistic Vesper spa r ro~~ regression models indicated that treatment (P= Pedestrian alone (on-trail1 9 25 (0 8 j M b 43 06 (3 9514 0001) and perpendicdar distance of the bird (Plt Dog on leash (on-trail) 10 13 (0 92)A 39 39 (4 56)A 0001) to the trail (for on-trail) or line of movement Dog alone (on-trail) 9 89 (1 85)A 35 41 (6 521A (for off-trail) were significant predictors of flush Pedestrian alone (off-trail) 16 95 10 87)B 51 49 15 44iA response (Figure 2) The shorter the perpendicular Dog on leash (otf-trail) 15 11 (0 89)B 52 23 (3 99iA distance of the robin to the trail or line of move- Dog alone (ott-trall) 10 87 (1 16)A 43 43 (5 91)A

ment the greater the probability that it wodd hlestern meadowlark

flush The area of influence was greater for off-trail 75 33 (6 55)AgcbPedestrian alone (on-trail) 30 63 (1 9 1 ) ~ ~

Dog on leash (on-trail) 28 21 11 52)A 65 68 (6 09jC than for on-trail treatments (Figure 2) However the

Dog alone l8 78 (234jB 9l 50 (7471BD area of influence did not differ between the pedes-

Pedestrian alone (ott-trail) 37 73 12 07)C 95 97 16 57)D trian-alone and dog-on-leash treatments either on-

Dog on leash (off-trail) 36 71 (1 50)C 102 29 (6 7310 or off-trail Date time of day and height of bird (if Dog alone off-trail) 33 50 (2 03jAC 88 75 (5 38)AD perched in tree) pre-flush were not significant pre-

dictors of whether a robin flushed (all Pgt005) a Distance between the actlvitv and bird when flushed When robins flushed mean flush distance dif- IJ Means with the rame letter within a column do not differ fered among treatnlents (F= 1792 plt0001

( P gt 0 05) Table 2) Flush distance was greater for off-trail treatments than for on-trail with the greatest flush

Among on-trail treatments flush distance was distance for the off-trail dog-on-leash treatment shorter for the dogalone treatment than either the Distance moved after flushing also differed among pedestrian-alone or dog-on-leash treatments which treatments (F3 129=350 P=0 017 Table 2) Dis-did not differ There were no differences in flush tance moved was greatest for the off-trail dog-on- distance among off-trail treatments For mead- leash treatment and shortest for the on-trail pedes- owlarks the distance moved after flushing dif- trian-alone treatment fered among treatments (F524- t=3993 p=0 002 Table 1) Distance moved Pedestrian alone (on-trail) Dog on leash (on-trail) was greater for a pedestri- an alone and a dog on leash when these activi- o 60l l l r ties occurred off-trail vs

P) 0 4 0on-trail On- and off-trail s 0 20dog-alone treatments did 5 OZ0 i i K inot differ Among on- = Oooo 5 10 15 20 25 30 o 000 5 10 15 20 25 30

trail treatments distance o moved differed only be- 2- Pedestrian alone (off-tra~l) Dog on leash (off-tra~l) tween the dog-on-leash 1 0 0

and dog-alone treatments 3 80

with the latter belng n 0 6 0

greater There were no differences in distance 0 40

moved among off-trail 0 20 o 20

treatments OoOO 5 10 5 20 25 30 OoOO 5 10 15 20 25 30

For each treatment flush distance was greater Meters from trail (for on-trail) or line of movement (for off-trail) for western Fiugre 2 Predicted probability of an American robin flushing to treatments in forests during than for vesper sparrows 1996 City of Boulder Open Space Boulder Colorado (dashed lines indicate 95 Cl)

Pedestrians and dogs Miller et al 125

requires an understanding of how wildlife responds to various forms of outdoor recreation and also the temporal and spatial context in which the activity occurs (Knight and Cole 1995)

Information on how wildlife reacts to hikers and dogs is limited although preliminary evidence sug-gests that presence of dogs increases the response For example mountain sheep (Ouis ccuzadensis MacArthur et al 19791982)golden plovers (Pluzli-nlis upricaria Yalden and Yalden 1990) and mar-mots (LMarmotcz marmotu Mainini et al 1993) exhibited a greater response when pedestrians were accompanied by a dog compared to solitary pedestrians

Location and frequency of recreational activities also can influence wildlife responses (Knight and Cole 1995) If animals perceive an activity as spa-tially predictable and nonthreatening they may habituate to that activity (Whittaker and Knight 1998) For example humans approaching from a parking area (an area with consistent hunlan use) elicited less of a response from mountain sheep than did humans approaching from over a ridge where human use was sporadic (MacArthur et al 1982)

Of the numerous studies on effects of recreation-al activities on wildlife most present information on flush distance (the distance between the activi-ty and the animal when it flushes) as the animal is approached directly by humans Although bird-watchers photographers and others do approach wildlife most recreationists do not go out of their way to do so Rather most recreationists such as hikers walking on trailsdo not commonly leave the trail To investigate this type of disturbancewe cor-related an animals flush response with its perpen-dicular distance to the trail or line of human move-ment With this informationwe were able to assess an area of influence for each treatment Area of influence was defined as the probability that an animal will flush or become alert (for mule deer only) at a given perpendicular distance from a trail or line of human movement The greater the area of influence the more disturbing the activity is to wildlife For example if the probability of flushing for a bird 30 m away from a trail is 040 to a pedes-trian accompanied by a dog and 070 to a pedestri-an alone then the area of influence is greater for the pedestrian alone

Our objective was to assess the area of influence around a lone pedestrian a pedestrian accompa-nied by a dog on leash and a dog aloneon and off trails For animals that flushedwe compared infor-

mation on flush distance (the distance between the activity and the animal when flushed) and distance moved to further assess the magnitude of distur-bance for each treatment Additionally for mule deer we compared illformation on alert distance (the distance between the activity and the deer when it became alert) among treatments In grass-lands we recorded responses of vesper sparrows and w-estern meadowlarks to all treatments In forests we recorded responses of American robins and mule deer to all treatments except the dog alone For each specieswe tested the null hypoth-esis that the area of influence and magnitude of dls-turbance did not differ between treatments

Methods and study area We conducted our study on 8000 ha of City of

Boulder Open Space property in and around the city of Boulder Colorado (40deg00N 105deg1845E) Elevation within the study area ranged from 1219 to 2438 m encompassing forest riparian shrub-land and grassland habitats Visitor use on City of Boulder Open Space is approximately 2 million vis-itsyear and is greatest during the spring followed by summer fall and winter (Zeller et al 1993) Recreational activities included hiking wildlife viewing exercising pets jogging mountain biking and horseback riding (hunting is not allowed)

We located study sites in pine forests and mixed-grass prairies Forests were dominated by pon-derosa pine (Pinus potzderosa) associated with shrubsgrasses and forbs Mixed-grass prairies con-tained a variety of tall mid-height and shortgrass species including little bluestem (Scbiznch-yriurn scopariurn) western wheatgrass (Agropj~ron smithiz3blue grama (Boz~telouagrcacilis) and side oats grama (Bouteloua curtipendz~la)

We conducted treatments on trails and for off-trail sites on areas gt400 m from trails Trails received frequent use whereas off-trail sites were used sporadically by recreationists We located all sites gt800 nl from urban developmentand gt400 m from physiographic features such as forest edge riparian areas and ridge lines Trail width was 125 k022 m (mean 1 SE) in the grasslands and 11-+ 020 m (mean 1 SE) in the forests

We collected data between 14 April and 20 July 1996 We rotated visits to on-trailand off-trailsites to avoid repeatedly sampling the same areas Birds and mule deer were not marked so we could not assure the same individuals were not mdtiply sampled

126 WWlfe Society Bulletin 200129(1) 124- 132

Grassland In grasslands we recorded responses of vesper

sparrows and western meadowlarks to 3 activities on- and off-trail 1) a pedestrian alone 2) a pedestri- an accompanied by a dog on leash and 3) a dog alone We selected these species because of their abundance on the study site and we were able to obtain adequate sample sizes for statistical compar- isons For treatments involving dogs we used either a 25-kg or a 40-kg dog Leash length was 18 m For on- and off-trail dog alone treatments the dog maintained an approximate distance of 20 m in front of the observer For dog-alone treatments we assumed that birds were responding to the dog only and not the observer In no case did the dogs attempt to chase birds

For on-trail treatments we detected individual birds on or near the trail ahead of us and proceed- ed along the center of the trail at approximately 15 msecond until the bird flushed or the observ- er had passed by eliciting no flush response At that time the observer stopped momentarily to record 1) flush response 2) the perpendicular distance between the bird and the trail 3) flush distance and 4) distance moved On off-trail sites we locat- ed birds on or near our line of movement and pro- ceeded parallel to the birds position so as to pass by at various distances (0 m to 200 m perpendicu- lar distance) After the bird flushed or the observer passed by eliciting no flush response we stopped momentarily to record the same information as that for on-trail treatments

Forest In forests we recorded responses of American

robins and mule deer both on- and off-trail to a pedestrian alone and a pedestrian accompanied by a dog on leash We selected these species because of their abundance on the study site and we were able to obtain adequate sample sizes for statistical comparisons Information for a dog alone was not recorded because we were unable to maintain an adequate distance behind the dog and still assume that robins or deer were responding only to the dog

We conducted treatments with robins and deer the same as in the grassland trials For robins we also measured (to the nearest 1 m) height above the ground (if perched in a tree) For deer we also recorded 1) alert response (ie lifted its head) 2) alert distance and 3) time elapsed from when a deer first exhibited a response until it resumed the

pre-disturbance behavior When group size was gt 1 we recorded information for the first deer to elicit a response We used a Lietz rangefinder (model 3390) to measure all distances to the nearest 1 m

Statistical analyses We used logistic regression (GENMOD proce-

dure SAS Institute Inc 1993) to determine whether flush response (and alert response for deer) of indi- vidual species was correlated with treatment per- pendicular distance to trail or line of movement date time of ampa) height of bird if perched in tree (for American robins) and group size and sex (for mule deer) For the animals that flushed we used analysis of variance (SAS Institute Inc 1988) to compare flush distance among treatments and also distance moved among treatments of individual species Because we attempted to simulate typical recreationist behavior (ie continuing to proceed along the trail or line of movement without stop- ping) many deer remained alert to our presence until we moved out of their sight Consequently mean and SE of time elapsed from when a deer first exhibited a response until it resumed the pre-disturbance activity could not be determined and we did not conduct statistical analysis comparing treatments For each grassland treatment we com- pared flush distance and also distance moved between vesper sparrows and western mead-owlarks using t-tests (SAS Institute Inc 1988) We used an a=005 for all analyses

Results Grassland

We conducted 462 and 393 trials for vesper spar- rows and western meadowlarks respectively For both species logistic regression models indicated that treatment (Plt0001) and perpendicular dis- tance of the bird (Plt0001) to a trail or line of movement (for off-trail) were significant predictors of flush response (Figure 1) The shorter the per- pendicular distance of a bird to the trail or line of movement the greater the probability that a bird would flush For both species the area of influence was greater for off-trail treatments than for on-trail treatments (Figure 1) For vesper sparrows on- and off-trail and also for western meadowlarks on-trail the dog-alone treatment resulted in a smaller area of influence than the pedestrian-alone or dog-on- leash treatments which did not differ from each other For western meadowlarks area of influence

Pedestrians and dogs Miller et al 127

(a) Pedestrian alone (on-trail) Dog on leash (on-trail) Dog alone (on-trail)

Pedestrian alone (off-trail) Dog on leash (off-trail) Dog alone (off-trail)

Pedestrian alone (on-trail) Dog on leash (on-trail) Dog alone (on-trail)(b)

rC

0 5 Pedestrian alone (off-trail) Dog on leash (off-trail)- Dog alone (off-trail)

~ ~~~~ ~ ~~

0 00

Meters from trail (for on-trail) or line of movement (for off-trail) Figure 1 Predicted probability of a vesper sparrow (a) and western meadowlark ibi flushing to treatments in grasslands during 1996 City of Boulder Open Space Boulder Colorado (dashed lines indicate 95 CIj

did not differ among off-trail treatments Date and time of day were not significant predictors of whether a bird would flush (Pgt005 for both species)

When vesper sparrows flushed mean flush dis-tance differed among treatments (F3269= 1175Plt 0001Table 1) Flush distance was greater for the off-trail pedestrian-alone and off-trail dog-on-leash treatments than for any other treatment Other

treatments did not differ For vesper sparrows dis-tance moved did not differ among treatments (Fj2= 146P=0204Table 1) however birds at off-trail sites tended to fly farther when compared to on-trail sites

When western meadowlarks flushed mean flush distance differed among treatments (Fj244=800 Plt0001Table 1) For each activityflush distance was greater for off-trail than on-trail treatments

128 Wildlife Sociep Bulletin 200129(1)124- 132

Table 1 Mean (SEI of flush distancea and distance moved for (t2298 Plt0005) and meadowlarks flew greater vesper sparrows and western meadowlarks in grasslands City distances once flushed (t340~10001) of Boulder Open Space Boulder Colorado 1996

Flush Distance FOfamp Treatment distance (m) moved ~ m We ran 228 trials for American robins Logistic Vesper spa r ro~~ regression models indicated that treatment (P= Pedestrian alone (on-trail1 9 25 (0 8 j M b 43 06 (3 9514 0001) and perpendicdar distance of the bird (Plt Dog on leash (on-trail) 10 13 (0 92)A 39 39 (4 56)A 0001) to the trail (for on-trail) or line of movement Dog alone (on-trail) 9 89 (1 85)A 35 41 (6 521A (for off-trail) were significant predictors of flush Pedestrian alone (off-trail) 16 95 10 87)B 51 49 15 44iA response (Figure 2) The shorter the perpendicular Dog on leash (otf-trail) 15 11 (0 89)B 52 23 (3 99iA distance of the robin to the trail or line of move- Dog alone (ott-trall) 10 87 (1 16)A 43 43 (5 91)A

ment the greater the probability that it wodd hlestern meadowlark

flush The area of influence was greater for off-trail 75 33 (6 55)AgcbPedestrian alone (on-trail) 30 63 (1 9 1 ) ~ ~

Dog on leash (on-trail) 28 21 11 52)A 65 68 (6 09jC than for on-trail treatments (Figure 2) However the

Dog alone l8 78 (234jB 9l 50 (7471BD area of influence did not differ between the pedes-

Pedestrian alone (ott-trail) 37 73 12 07)C 95 97 16 57)D trian-alone and dog-on-leash treatments either on-

Dog on leash (off-trail) 36 71 (1 50)C 102 29 (6 7310 or off-trail Date time of day and height of bird (if Dog alone off-trail) 33 50 (2 03jAC 88 75 (5 38)AD perched in tree) pre-flush were not significant pre-

dictors of whether a robin flushed (all Pgt005) a Distance between the actlvitv and bird when flushed When robins flushed mean flush distance dif- IJ Means with the rame letter within a column do not differ fered among treatnlents (F= 1792 plt0001

( P gt 0 05) Table 2) Flush distance was greater for off-trail treatments than for on-trail with the greatest flush

Among on-trail treatments flush distance was distance for the off-trail dog-on-leash treatment shorter for the dogalone treatment than either the Distance moved after flushing also differed among pedestrian-alone or dog-on-leash treatments which treatments (F3 129=350 P=0 017 Table 2) Dis-did not differ There were no differences in flush tance moved was greatest for the off-trail dog-on- distance among off-trail treatments For mead- leash treatment and shortest for the on-trail pedes- owlarks the distance moved after flushing dif- trian-alone treatment fered among treatments (F524- t=3993 p=0 002 Table 1) Distance moved Pedestrian alone (on-trail) Dog on leash (on-trail) was greater for a pedestri- an alone and a dog on leash when these activi- o 60l l l r ties occurred off-trail vs

P) 0 4 0on-trail On- and off-trail s 0 20dog-alone treatments did 5 OZ0 i i K inot differ Among on- = Oooo 5 10 15 20 25 30 o 000 5 10 15 20 25 30

trail treatments distance o moved differed only be- 2- Pedestrian alone (off-tra~l) Dog on leash (off-tra~l) tween the dog-on-leash 1 0 0

and dog-alone treatments 3 80

with the latter belng n 0 6 0

greater There were no differences in distance 0 40

moved among off-trail 0 20 o 20

treatments OoOO 5 10 5 20 25 30 OoOO 5 10 15 20 25 30

For each treatment flush distance was greater Meters from trail (for on-trail) or line of movement (for off-trail) for western Fiugre 2 Predicted probability of an American robin flushing to treatments in forests during than for vesper sparrows 1996 City of Boulder Open Space Boulder Colorado (dashed lines indicate 95 Cl)

126 WWlfe Society Bulletin 200129(1) 124- 132

Grassland In grasslands we recorded responses of vesper

sparrows and western meadowlarks to 3 activities on- and off-trail 1) a pedestrian alone 2) a pedestri- an accompanied by a dog on leash and 3) a dog alone We selected these species because of their abundance on the study site and we were able to obtain adequate sample sizes for statistical compar- isons For treatments involving dogs we used either a 25-kg or a 40-kg dog Leash length was 18 m For on- and off-trail dog alone treatments the dog maintained an approximate distance of 20 m in front of the observer For dog-alone treatments we assumed that birds were responding to the dog only and not the observer In no case did the dogs attempt to chase birds

For on-trail treatments we detected individual birds on or near the trail ahead of us and proceed- ed along the center of the trail at approximately 15 msecond until the bird flushed or the observ- er had passed by eliciting no flush response At that time the observer stopped momentarily to record 1) flush response 2) the perpendicular distance between the bird and the trail 3) flush distance and 4) distance moved On off-trail sites we locat- ed birds on or near our line of movement and pro- ceeded parallel to the birds position so as to pass by at various distances (0 m to 200 m perpendicu- lar distance) After the bird flushed or the observer passed by eliciting no flush response we stopped momentarily to record the same information as that for on-trail treatments

Forest In forests we recorded responses of American

robins and mule deer both on- and off-trail to a pedestrian alone and a pedestrian accompanied by a dog on leash We selected these species because of their abundance on the study site and we were able to obtain adequate sample sizes for statistical comparisons Information for a dog alone was not recorded because we were unable to maintain an adequate distance behind the dog and still assume that robins or deer were responding only to the dog

We conducted treatments with robins and deer the same as in the grassland trials For robins we also measured (to the nearest 1 m) height above the ground (if perched in a tree) For deer we also recorded 1) alert response (ie lifted its head) 2) alert distance and 3) time elapsed from when a deer first exhibited a response until it resumed the

pre-disturbance behavior When group size was gt 1 we recorded information for the first deer to elicit a response We used a Lietz rangefinder (model 3390) to measure all distances to the nearest 1 m

Statistical analyses We used logistic regression (GENMOD proce-

dure SAS Institute Inc 1993) to determine whether flush response (and alert response for deer) of indi- vidual species was correlated with treatment per- pendicular distance to trail or line of movement date time of ampa) height of bird if perched in tree (for American robins) and group size and sex (for mule deer) For the animals that flushed we used analysis of variance (SAS Institute Inc 1988) to compare flush distance among treatments and also distance moved among treatments of individual species Because we attempted to simulate typical recreationist behavior (ie continuing to proceed along the trail or line of movement without stop- ping) many deer remained alert to our presence until we moved out of their sight Consequently mean and SE of time elapsed from when a deer first exhibited a response until it resumed the pre-disturbance activity could not be determined and we did not conduct statistical analysis comparing treatments For each grassland treatment we com- pared flush distance and also distance moved between vesper sparrows and western mead-owlarks using t-tests (SAS Institute Inc 1988) We used an a=005 for all analyses

Results Grassland

We conducted 462 and 393 trials for vesper spar- rows and western meadowlarks respectively For both species logistic regression models indicated that treatment (Plt0001) and perpendicular dis- tance of the bird (Plt0001) to a trail or line of movement (for off-trail) were significant predictors of flush response (Figure 1) The shorter the per- pendicular distance of a bird to the trail or line of movement the greater the probability that a bird would flush For both species the area of influence was greater for off-trail treatments than for on-trail treatments (Figure 1) For vesper sparrows on- and off-trail and also for western meadowlarks on-trail the dog-alone treatment resulted in a smaller area of influence than the pedestrian-alone or dog-on- leash treatments which did not differ from each other For western meadowlarks area of influence

Pedestrians and dogs Miller et al 127

(a) Pedestrian alone (on-trail) Dog on leash (on-trail) Dog alone (on-trail)

Pedestrian alone (off-trail) Dog on leash (off-trail) Dog alone (off-trail)

Pedestrian alone (on-trail) Dog on leash (on-trail) Dog alone (on-trail)(b)

rC

0 5 Pedestrian alone (off-trail) Dog on leash (off-trail)- Dog alone (off-trail)

~ ~~~~ ~ ~~

0 00

Meters from trail (for on-trail) or line of movement (for off-trail) Figure 1 Predicted probability of a vesper sparrow (a) and western meadowlark ibi flushing to treatments in grasslands during 1996 City of Boulder Open Space Boulder Colorado (dashed lines indicate 95 CIj

did not differ among off-trail treatments Date and time of day were not significant predictors of whether a bird would flush (Pgt005 for both species)

When vesper sparrows flushed mean flush dis-tance differed among treatments (F3269= 1175Plt 0001Table 1) Flush distance was greater for the off-trail pedestrian-alone and off-trail dog-on-leash treatments than for any other treatment Other

treatments did not differ For vesper sparrows dis-tance moved did not differ among treatments (Fj2= 146P=0204Table 1) however birds at off-trail sites tended to fly farther when compared to on-trail sites

When western meadowlarks flushed mean flush distance differed among treatments (Fj244=800 Plt0001Table 1) For each activityflush distance was greater for off-trail than on-trail treatments

128 Wildlife Sociep Bulletin 200129(1)124- 132

Table 1 Mean (SEI of flush distancea and distance moved for (t2298 Plt0005) and meadowlarks flew greater vesper sparrows and western meadowlarks in grasslands City distances once flushed (t340~10001) of Boulder Open Space Boulder Colorado 1996

Flush Distance FOfamp Treatment distance (m) moved ~ m We ran 228 trials for American robins Logistic Vesper spa r ro~~ regression models indicated that treatment (P= Pedestrian alone (on-trail1 9 25 (0 8 j M b 43 06 (3 9514 0001) and perpendicdar distance of the bird (Plt Dog on leash (on-trail) 10 13 (0 92)A 39 39 (4 56)A 0001) to the trail (for on-trail) or line of movement Dog alone (on-trail) 9 89 (1 85)A 35 41 (6 521A (for off-trail) were significant predictors of flush Pedestrian alone (off-trail) 16 95 10 87)B 51 49 15 44iA response (Figure 2) The shorter the perpendicular Dog on leash (otf-trail) 15 11 (0 89)B 52 23 (3 99iA distance of the robin to the trail or line of move- Dog alone (ott-trall) 10 87 (1 16)A 43 43 (5 91)A

ment the greater the probability that it wodd hlestern meadowlark

flush The area of influence was greater for off-trail 75 33 (6 55)AgcbPedestrian alone (on-trail) 30 63 (1 9 1 ) ~ ~

Dog on leash (on-trail) 28 21 11 52)A 65 68 (6 09jC than for on-trail treatments (Figure 2) However the

Dog alone l8 78 (234jB 9l 50 (7471BD area of influence did not differ between the pedes-

Pedestrian alone (ott-trail) 37 73 12 07)C 95 97 16 57)D trian-alone and dog-on-leash treatments either on-

Dog on leash (off-trail) 36 71 (1 50)C 102 29 (6 7310 or off-trail Date time of day and height of bird (if Dog alone off-trail) 33 50 (2 03jAC 88 75 (5 38)AD perched in tree) pre-flush were not significant pre-

dictors of whether a robin flushed (all Pgt005) a Distance between the actlvitv and bird when flushed When robins flushed mean flush distance dif- IJ Means with the rame letter within a column do not differ fered among treatnlents (F= 1792 plt0001

( P gt 0 05) Table 2) Flush distance was greater for off-trail treatments than for on-trail with the greatest flush

Among on-trail treatments flush distance was distance for the off-trail dog-on-leash treatment shorter for the dogalone treatment than either the Distance moved after flushing also differed among pedestrian-alone or dog-on-leash treatments which treatments (F3 129=350 P=0 017 Table 2) Dis-did not differ There were no differences in flush tance moved was greatest for the off-trail dog-on- distance among off-trail treatments For mead- leash treatment and shortest for the on-trail pedes- owlarks the distance moved after flushing dif- trian-alone treatment fered among treatments (F524- t=3993 p=0 002 Table 1) Distance moved Pedestrian alone (on-trail) Dog on leash (on-trail) was greater for a pedestri- an alone and a dog on leash when these activi- o 60l l l r ties occurred off-trail vs

P) 0 4 0on-trail On- and off-trail s 0 20dog-alone treatments did 5 OZ0 i i K inot differ Among on- = Oooo 5 10 15 20 25 30 o 000 5 10 15 20 25 30

trail treatments distance o moved differed only be- 2- Pedestrian alone (off-tra~l) Dog on leash (off-tra~l) tween the dog-on-leash 1 0 0

and dog-alone treatments 3 80

with the latter belng n 0 6 0

greater There were no differences in distance 0 40

moved among off-trail 0 20 o 20

treatments OoOO 5 10 5 20 25 30 OoOO 5 10 15 20 25 30

For each treatment flush distance was greater Meters from trail (for on-trail) or line of movement (for off-trail) for western Fiugre 2 Predicted probability of an American robin flushing to treatments in forests during than for vesper sparrows 1996 City of Boulder Open Space Boulder Colorado (dashed lines indicate 95 Cl)

Pedestrians and dogs Miller et al 127

(a) Pedestrian alone (on-trail) Dog on leash (on-trail) Dog alone (on-trail)

Pedestrian alone (off-trail) Dog on leash (off-trail) Dog alone (off-trail)

Pedestrian alone (on-trail) Dog on leash (on-trail) Dog alone (on-trail)(b)

rC

0 5 Pedestrian alone (off-trail) Dog on leash (off-trail)- Dog alone (off-trail)

~ ~~~~ ~ ~~

0 00

Meters from trail (for on-trail) or line of movement (for off-trail) Figure 1 Predicted probability of a vesper sparrow (a) and western meadowlark ibi flushing to treatments in grasslands during 1996 City of Boulder Open Space Boulder Colorado (dashed lines indicate 95 CIj

did not differ among off-trail treatments Date and time of day were not significant predictors of whether a bird would flush (Pgt005 for both species)

When vesper sparrows flushed mean flush dis-tance differed among treatments (F3269= 1175Plt 0001Table 1) Flush distance was greater for the off-trail pedestrian-alone and off-trail dog-on-leash treatments than for any other treatment Other

treatments did not differ For vesper sparrows dis-tance moved did not differ among treatments (Fj2= 146P=0204Table 1) however birds at off-trail sites tended to fly farther when compared to on-trail sites

When western meadowlarks flushed mean flush distance differed among treatments (Fj244=800 Plt0001Table 1) For each activityflush distance was greater for off-trail than on-trail treatments

128 Wildlife Sociep Bulletin 200129(1)124- 132

Table 1 Mean (SEI of flush distancea and distance moved for (t2298 Plt0005) and meadowlarks flew greater vesper sparrows and western meadowlarks in grasslands City distances once flushed (t340~10001) of Boulder Open Space Boulder Colorado 1996

Flush Distance FOfamp Treatment distance (m) moved ~ m We ran 228 trials for American robins Logistic Vesper spa r ro~~ regression models indicated that treatment (P= Pedestrian alone (on-trail1 9 25 (0 8 j M b 43 06 (3 9514 0001) and perpendicdar distance of the bird (Plt Dog on leash (on-trail) 10 13 (0 92)A 39 39 (4 56)A 0001) to the trail (for on-trail) or line of movement Dog alone (on-trail) 9 89 (1 85)A 35 41 (6 521A (for off-trail) were significant predictors of flush Pedestrian alone (off-trail) 16 95 10 87)B 51 49 15 44iA response (Figure 2) The shorter the perpendicular Dog on leash (otf-trail) 15 11 (0 89)B 52 23 (3 99iA distance of the robin to the trail or line of move- Dog alone (ott-trall) 10 87 (1 16)A 43 43 (5 91)A

ment the greater the probability that it wodd hlestern meadowlark

flush The area of influence was greater for off-trail 75 33 (6 55)AgcbPedestrian alone (on-trail) 30 63 (1 9 1 ) ~ ~

Dog on leash (on-trail) 28 21 11 52)A 65 68 (6 09jC than for on-trail treatments (Figure 2) However the

Dog alone l8 78 (234jB 9l 50 (7471BD area of influence did not differ between the pedes-

Pedestrian alone (ott-trail) 37 73 12 07)C 95 97 16 57)D trian-alone and dog-on-leash treatments either on-

Dog on leash (off-trail) 36 71 (1 50)C 102 29 (6 7310 or off-trail Date time of day and height of bird (if Dog alone off-trail) 33 50 (2 03jAC 88 75 (5 38)AD perched in tree) pre-flush were not significant pre-

dictors of whether a robin flushed (all Pgt005) a Distance between the actlvitv and bird when flushed When robins flushed mean flush distance dif- IJ Means with the rame letter within a column do not differ fered among treatnlents (F= 1792 plt0001

( P gt 0 05) Table 2) Flush distance was greater for off-trail treatments than for on-trail with the greatest flush

Among on-trail treatments flush distance was distance for the off-trail dog-on-leash treatment shorter for the dogalone treatment than either the Distance moved after flushing also differed among pedestrian-alone or dog-on-leash treatments which treatments (F3 129=350 P=0 017 Table 2) Dis-did not differ There were no differences in flush tance moved was greatest for the off-trail dog-on- distance among off-trail treatments For mead- leash treatment and shortest for the on-trail pedes- owlarks the distance moved after flushing dif- trian-alone treatment fered among treatments (F524- t=3993 p=0 002 Table 1) Distance moved Pedestrian alone (on-trail) Dog on leash (on-trail) was greater for a pedestri- an alone and a dog on leash when these activi- o 60l l l r ties occurred off-trail vs

P) 0 4 0on-trail On- and off-trail s 0 20dog-alone treatments did 5 OZ0 i i K inot differ Among on- = Oooo 5 10 15 20 25 30 o 000 5 10 15 20 25 30

trail treatments distance o moved differed only be- 2- Pedestrian alone (off-tra~l) Dog on leash (off-tra~l) tween the dog-on-leash 1 0 0

and dog-alone treatments 3 80

with the latter belng n 0 6 0

greater There were no differences in distance 0 40

moved among off-trail 0 20 o 20

treatments OoOO 5 10 5 20 25 30 OoOO 5 10 15 20 25 30

For each treatment flush distance was greater Meters from trail (for on-trail) or line of movement (for off-trail) for western Fiugre 2 Predicted probability of an American robin flushing to treatments in forests during than for vesper sparrows 1996 City of Boulder Open Space Boulder Colorado (dashed lines indicate 95 Cl)

128 Wildlife Sociep Bulletin 200129(1)124- 132

Table 1 Mean (SEI of flush distancea and distance moved for (t2298 Plt0005) and meadowlarks flew greater vesper sparrows and western meadowlarks in grasslands City distances once flushed (t340~10001) of Boulder Open Space Boulder Colorado 1996

Flush Distance FOfamp Treatment distance (m) moved ~ m We ran 228 trials for American robins Logistic Vesper spa r ro~~ regression models indicated that treatment (P= Pedestrian alone (on-trail1 9 25 (0 8 j M b 43 06 (3 9514 0001) and perpendicdar distance of the bird (Plt Dog on leash (on-trail) 10 13 (0 92)A 39 39 (4 56)A 0001) to the trail (for on-trail) or line of movement Dog alone (on-trail) 9 89 (1 85)A 35 41 (6 521A (for off-trail) were significant predictors of flush Pedestrian alone (off-trail) 16 95 10 87)B 51 49 15 44iA response (Figure 2) The shorter the perpendicular Dog on leash (otf-trail) 15 11 (0 89)B 52 23 (3 99iA distance of the robin to the trail or line of move- Dog alone (ott-trall) 10 87 (1 16)A 43 43 (5 91)A

ment the greater the probability that it wodd hlestern meadowlark

flush The area of influence was greater for off-trail 75 33 (6 55)AgcbPedestrian alone (on-trail) 30 63 (1 9 1 ) ~ ~

Dog on leash (on-trail) 28 21 11 52)A 65 68 (6 09jC than for on-trail treatments (Figure 2) However the

Dog alone l8 78 (234jB 9l 50 (7471BD area of influence did not differ between the pedes-

Pedestrian alone (ott-trail) 37 73 12 07)C 95 97 16 57)D trian-alone and dog-on-leash treatments either on-

Dog on leash (off-trail) 36 71 (1 50)C 102 29 (6 7310 or off-trail Date time of day and height of bird (if Dog alone off-trail) 33 50 (2 03jAC 88 75 (5 38)AD perched in tree) pre-flush were not significant pre-

dictors of whether a robin flushed (all Pgt005) a Distance between the actlvitv and bird when flushed When robins flushed mean flush distance dif- IJ Means with the rame letter within a column do not differ fered among treatnlents (F= 1792 plt0001

( P gt 0 05) Table 2) Flush distance was greater for off-trail treatments than for on-trail with the greatest flush

Among on-trail treatments flush distance was distance for the off-trail dog-on-leash treatment shorter for the dogalone treatment than either the Distance moved after flushing also differed among pedestrian-alone or dog-on-leash treatments which treatments (F3 129=350 P=0 017 Table 2) Dis-did not differ There were no differences in flush tance moved was greatest for the off-trail dog-on- distance among off-trail treatments For mead- leash treatment and shortest for the on-trail pedes- owlarks the distance moved after flushing dif- trian-alone treatment fered among treatments (F524- t=3993 p=0 002 Table 1) Distance moved Pedestrian alone (on-trail) Dog on leash (on-trail) was greater for a pedestri- an alone and a dog on leash when these activi- o 60l l l r ties occurred off-trail vs

P) 0 4 0on-trail On- and off-trail s 0 20dog-alone treatments did 5 OZ0 i i K inot differ Among on- = Oooo 5 10 15 20 25 30 o 000 5 10 15 20 25 30

trail treatments distance o moved differed only be- 2- Pedestrian alone (off-tra~l) Dog on leash (off-tra~l) tween the dog-on-leash 1 0 0

and dog-alone treatments 3 80

with the latter belng n 0 6 0

greater There were no differences in distance 0 40

moved among off-trail 0 20 o 20

treatments OoOO 5 10 5 20 25 30 OoOO 5 10 15 20 25 30

For each treatment flush distance was greater Meters from trail (for on-trail) or line of movement (for off-trail) for western Fiugre 2 Predicted probability of an American robin flushing to treatments in forests during than for vesper sparrows 1996 City of Boulder Open Space Boulder Colorado (dashed lines indicate 95 Cl)

Pedestrians and dogs Miller et al 129

Table 2 Mean ISE) of flush distanceQnd distance moved for area of influence was greater for off-trailtreatments American robins in Forests City of Boulder Open Space Bod- than for on-trail (Figure 3) For both on- and off-der Colorado 1996 trail area of influence was greater when a dog was

Flush Distance present Deer group sizesexdate and time of day

Treatment distance (m) of lush imr were not significant predictors of whether a deer

Pedestrian alone ion-trail) 961 ( 0 6 3 ) ~ ~1497 (219)At would flush (all Pgt005)

Dog on leash on-trail) 982 05jA 2079 (209AB When deer flushed mean flush distance differed

Pedestrian alone (oii-trail) 1374 t1 08B 1731 (185)AB (F342= PltOOOO1l D~~on leashiofftraill 1627 ( 0 6 0 ~ 2349 ( 2 0 5 ) ~ 3) Flush distance was greater for the off-trail dog-

on-leash treatment than any other Because many of V i s t a n c e between the activity and bird when flushed the deer that flushed moved out of sight for the off-

Means with the same letter within a column do not difier trail dog-on-leaslltreatmentwe could not calculate ( P gt 0051 mean and SE of distance moved for this treatment

Therefore we did not include the off-trail dog-on-We ran 88trials for mule deer Logistic regression leash treatment in statistical comparisons of dis-

models indicated that treatment (P=0003) and tance moved among treatments When comparing perpendicular distance of the deer (P=0002) to the other treatments distance moved differed the trail or line of movement (when off-rail) were between treatments (F230=780P=0002Table3) significant predictors of alert response (Figure 3) Distance moved was greater for the off-trail pedes-For on-trail treatments the shorter the perpendicu- trian-alone treatment than the on-trail treatments lar distance of deer to trail the greater the proba- which did not differ bility that it would become alert The area of influ-ence was greatest for off-trailtreatments where the deer became alert regardless of activity type or Discussion their perpendicular distance to the line of move- Wildlife may exhibit diverse responses to arious ment (Figure 3) On-trail the dog-on-leash treat- types of recreational activities and may be influ-ment resulted in a greater area of influence than enced by the frequency and spatial context in the pedestrian-alone treatment Deer group size which the activity occurs (Knight and Cole 1995) sex date and time of day were not significant In general for vesper sparrows and western predictors of whether a deer would become alert meadowlarks the flush distance and distance (all Pgt005) When deer did become alert mean moved was shortest and the area of influence was alert distance differed among treatments (F = smallest for dog-alone treatments and greatest 797 Plt0001 Table 3) When comparing each when a pedestrian was present Because dogs activit) individually there were no differences in closely resemble coyotes (Canis l~zt~c~tzs)and foxes alert distance whether the activity occurred on- or (VtlZpesfulria) and because these species are hpi-off-trail Howeverwithin on-or off-trailtreatments cally not considered significant predators on song-alert distance was greater

when a dog was presellt Table 3 Mean and SE of alert distancea tlush distanceh and distance movedc tor mule deer Logistic regression mo- in forests City of Boulder Open Space Boulder Colorado 1996

dels indicated that treat-ment (Plt0001) and per- Alert Flush D~stance

pendicular distance of the Treatment d~stance(mi d~stance(mi moved (nil

deer ( p = o 001) to the Pedestr~analone on-tra~li 45 55 (12 75)Ad 33 50 (0 50iAd 31 50 (1 50)Ad

trail (for on-trail) and line Dog odeash 85 37 18 13)B C 48 50 I 751A 33 89 1596rA

of movement (for off-trail) were significant predic-tors of flush response (Figure 3) The closer the deer was to the trail or line of movement the greater the probability that it would flush The

Pedestr~analone (otf-tra~l) 66 77 (4 34IAB 34 19 14 63)A 77 0 19 61)B

Dog on leash (otf-tra~l) 100 60 17 811C 81 92 1785)B rgt76 - gt300e

V i s t a n c e between the activity and deer when it became alert

I Distance between the activitv and deer when it flushed

Mean and SE coultl not be determined because some cleer moved out of view for the

dog on leash treatment therefore this treatment was not included in the analys~s

Means lvith the same letter do not differ P gt 005)

Indicates range of distance moved beiore deer moved out of view

1 30 uiildlijk Society Rzrlletin 2 0 0 129(1) 124 - 132

Pedestrian alone (on-tra~l) Dog on leash (on-trail) a pedestrian 1s the addi- 1 00 tive Factor Thls is further

o 80 supported by the fact that the area of influence did

0 60 not differ between the

0 40 0 40 pedestrian-alone and dog- o 20 o 20 on-leash treatments for

0000 20 40 60 80 100 o 00

0 20 40 60 80 100 American robins in forest either on- or

the off--3 Pedestrian-alone (off-trail) Dog on leash (off-trail) trail

For all species in our study area of influence flush distance and dis-tance moved were almost always greater when activities occurred off-trail

0 00 o 00 versus when the same 0 20 40 60 60 100 0 20 40 60 80 100

activih occurred on-trail Recreational use occurred

Pedestrian-alone (on-trail) Dog on leash (on-trail) on our off-trail study sites but was sporadic con-versely recreational use on trails was common (City of Boulder Open Space 1996) Because recreational activities oc- curring on-trail were fre-quent and spatially pre-

Pedestrian alone (off-trail) Dog on leash (off-trail) dictable animals had likely habituated to activi- ty in these locations Off- trail recreation however was infrequent and spa-tially unpredictable Thus animals were not accus-tomed to activity in these areas resulting in the

Meters from trail (for on-trail) or line of movement (for off-trail) greater area of influence

Fiugre 2 Predicted probability of an American robin flushing to treatments in forests during flush distance and dis-1996 City ol Boulder Open Space Boulder Colorado (dashed lines indicate 95 CI) tance moved In Switzer-

land a study of marmots revealed similar results

birds (Leach and Frazier 1953Andelt et al 1987) (Mainini et al 1993) They found that marmots these bird species may not have perceived dogs as exhibited the greatest response to hikers when hik- an important threat Alternatively dogs may pose a ers strayed away from trails Cooke (1 980) Yalden different kind of threat than a pedestrian and birds andYalden (1989) Burger and Gochfeld (1991) and may hold their position until the last moment Kenny and Knight (1992) showed that in areas attempting to remain undetected Because the area where human activity was common and frequent of influence was generally the smallest for the dog- birds were less disturbed than those in areas where alone treatments and because there were no signif- humans were uncommon Likewise Schultz and icant differences between the pedestrian-alone and Bailey (1978) MacArthur et al (1982) and Hamr dog-on-leash treatments it appears that presence of (1988) found that large mammals exhibited the

Pedestrians and dogs Miller et al 131

greatest response when human activity was spatial- Temple 1995) Because off-trailtreatments resulted 1y unpredictable in the greatest area of influence for all wildlife in

Unlike the responses of bird species in our study this study recreational use could be restricted mule deer exhibited the greatest response when a (through education and enforcement) to trails as a dog was present Similar to our results MacArthur way to reduce impacts However because negative et al (1979 1982) and Mainini et al (1993) found impacts occur even from on-trail use number and that lno~lntainsheep and nyarmots respectively spatial arrallgement of trails must be considered in exhibited heightened responses when dogs were conservation planning Furthermore kcause QPe

present ~ l thoughCit)of Boulder Open Space reg- of recreational activity influenced the magnitude of

ulations recluire that dogs be under control wildlife response managers could restrict certain

there were no leash laws 011 our study sites and recreatiollal activities such as prohibiting dogs in

dogs are known to brass alld attack deer (person- some areas or requiring dogs to be leashed Pdrti-

obsemtion) Because dogs can kill deer tiolling the landscape into recreation zones allow-

1953br ick 1969LOWryand Mcampthur 1978) arid ing certain activities in some zones while restrict-

because canicls have on deer tllroLlghollt ing them in others may aid in reducing conflicts

their evolutionary history we assume that deer with sensitive species

have become sensitized to the presence of dogs People are often not aware of h o ~ vtheir activities

explailling the greater reaction when a pedestrian affect wildlife even if the see animals respond to

was accompanied by a dog their actions (Stalmaster and Kaiser 1998) Even F ~ ) ~the species measured ill our studythe area of tllough the dog-alone treatment resulted in the

influence was slnaller whetl treatments occurred smallest area of influence for grassland birds in our

on-trail than off-trail However all species appeared study area of influence will increase if recreation-

to have a threshold of tolerance to disturbance ists alloartheir dogs to roatn amrayfrom a trail Addi-tionally in our study we did not stop and view thcbased on distance with a greater flush response subjects for extended periods of time or attempt to(and alert response for mule deer) when wildlife move toward them Behaviors of this kind are com-

were close to trails An earlier study on the same moll among nature viewers and could lead to ele-

area revealed a positive correlation between abun-vated wildlife responses (mein 1993)

dance of some bird species nest occurrence and Recreationists are more likely to support restric-

nest success with distance from trails r miller et al tions if they understand how wildlife will benefit

1998) The authors felt that this correlation was in (Purcly et al 1987Harris et al 1995) By emphasizingpart a result of recreational actirih and the associ- how human affect wildliie people can ass-ated disturbance Even though the area of influ- ciate their actions or harming encc for all species was sn~alleron-trail versus off- animal populations and begin to develop a conser-a-trail on-trail activities may still constitute an Such an ethic call minilllize important source of disturbance Thus our results wildlife-human conflicts occurring in llatural suggest that human activities may displace wildlife areas ( ~ ~ i ~ h ~and ~~~~l~ 1995) Klein(1993) found and reduce fitness in local wildlife populations A-hat visitors who spoke to wildlife refilge personnel mentioned earlieroff-trailrecreational use was spo- less likely to disturb wildlife than radic However should recreational use away from ho did not llus effectivevisitoreducation can aid trails increase displacelllent of wildlife may ~llti- in developing a collsemation ethic T13rough edllca-mately result Experiments conducted in forested tion land managers can inform recreatiollists how areas of Kyoming without trails Support this con- their activities affectwildlife and how the- can mod-clusion (Gutzwiller et al 1994 Riffell et al 1996 ify their behavior to minimize impacts Gutzwiller et al 1997) showing that recreational activities away from trails resulted in altered behav- Acknoudedgments We thank Madison ancl Bo for

ior and displacement of birds their assistance in the field We are grateful to the City of Boulder Open Space Department for finan-cial support

Management implications Land managers can use spatial and behavioral Literature cited

restrictions in management to L k A l gt ~ I ~ VFJ ( KIFF F K U O U T I O A ~ I )K C ~ I ) Ulili 19X7 Varj-istence of wildlife and recreationists (Knight and ation in coote diets associatecl with season and successional

132 WtXdlt~2Society Bulletin 200123(1) 124- 132

RA GEISSAZID 1982 Cardiac and 2-3-27 behavioral responses of mountain sheep to human distur-

BARICK 1969 Deer predation in North Carolina and other bance Journal of Wildlife Management 46 35 1-358

changes in vegetation Journal of Wildlife Management 51 RL~C~RTHLIII R HJOHYSTOU

F B southeastern states Pages 25-31 it2 White-tailed deer in the ~ZIINISIBI NELIFWL l I ~ ( o I SAUD 11 1993 Behaviour of marmots southern forest habitat Proceeding of a symposi~un United lfnnnota nlarmota under the influence of different hiking States Forest Service Southern Forest Experiment Station activities Biological Consen-ation 64 161-164 Nacogdoches Texas USA MILLER AVLI C K MIILER S G R L K~IGHT 1998 Influence of

BOWERSR R 1953 The free-nmnning dog menace Virginia recreational trails on breeding bird communities Ecological Wildlife 14 5- Applications 8 162-169

BOILE S A AND I B SAISON 1985 Effects of nonconsumptive PLRDIK G G R GOFIDJDLCKERGA P~VEK~UTY ~UL~ NACol-recreation on wildlife a review Wildlife Society Bulletin 13 x~111 198 A guide to managing 11umtn activity 011 Nation-110-116 al Wildlife Refuges Human Dimensions Resources Unit

BURGER M GOCHFELD Department of Natural Resources Cornell Lrniversih- Ithaca JAUI) 1991 Human distance and birds tol- erance and response distances of resident and migrant New York ITSA species in India Environmental Consenration 18 158-165 RIFFELL AUD ANI)ERSO~ DoesS K K J GITZ~TLLER S 1 1996

C m OF B ~ L L D E ROPE^ SFACE 1996 North Boulder Valley inven- repeated human intrusion cause c~~mulative declines in avian tory report City of Boulder Open Space Department Boul- richness and abundance Ecological Applications 6 492-505 der Colorado USA SAS IU5TlTl TE Isc 1988 SASSTAT I-sers Guide Release 603

Coo~cAS 1980 Observations on how close certain passerine Edition SAS Institute Inc Cay North Carolina USA species will tolerate an approaching human in rural and sub- SAS Iha~rrr rh IYC 1993 SAS Technical Report p-243 SASSTAT urban areas Biological Conservation 1885-88 Software the GENRlOD procedure Release 609 SAS Insti-

GLTZWTLLERK J E A KROESES H ~VUERSOU C A WILKINS AND tute Inc Cay North Carolina USA 199 Does human intrusion alter the seasonal timing of avian song during breeding periods Auk 114 55-65

Gri ZWIILER K L CLE~IEUTSKJ RT W I E D E ~ ~ N N ANI) 5 HAuDIR- sos 1994 Effects of human intrusions on song occurrence and singing consistency in subalpine birds Auk 11 1 28-37

H ~ M K 1988 Disturbance behaviour of chamois in an alpine J tourist area of Austria Mountain Research and Development 865-73

HARRIS AUD WW SHAY L K R R K R ~ L ~ S ~ M N 1995 Human atti- tudes and mountain sheep in a wilderness setting Wildlife Society Bulletin 2366-2

KEXWSA R L KNIGHT 1992 Flight distances of black- billed magpies in different regimes of human densih and persecution Condor 94 545- 547

KLEIN 1993 Waterbird behavioral responses to human dis- M L turbances Wildlife Society Bulletin 21 31 -39

KSIGHTR LAYD D N COIE 1995 Factors that influence wildlife responses to recreationists Pages 1-9 in Wildlife and recreationists coexistence through research and manage- ment R L Knight and K J Gutnl-iller editors Island Cov- elo California USA

KNIGHT 1995 Wildlife and recreation- R L~ N I )K J GL TZWILLER

ists coexistence through research and management R L Knight and K Gutzwiller editors Island Covelo California USA

KYIGHT 1995 Wildlife and recreationists R L A ~ L )SATEMPLE coexistence through management Pages 32-333 in Wildlife and recreationists coexistence through research and management R L Knight and K J Gutn7iller editors Island Corelo California USA

LWCHH R IUD W H FMIER 1953 A study of the possible extent of predation on heavy concentrations of valley quail with special reference to the bobcat California Fish and Game 39 52-538

LOWRY 1978 Domestic dogs as pred- DAAND K L ampI(-ARTHUR ators on deer Wildlife Society Bulletin 6 38-39

RIACARTHLR GEIST Factors influ- RA R H JOHNSTOSA~DV 19-9 encing heart rate in free-ranging bighorn sheep a physiolog-ical approach to the study of wildlife harassment Canadian Journal of Zoolog)- 572010-2021

SCHLLTZR DID JA BAILII 198 R~S~OIISCSof national park elk to human activity Journal of Wildlife lanagenlent i2 91-100

STampIJI~STFR J L KAISER Effect of recreational Y1 AD 1998 activity on wintering bald eagles Wiltllife Monograph 13-

WH~-laakRD ampnu RL KUIGHT 1998 Understanding wildlife responses to humans Wildlife Society Bulletin 26 312-31

YALDE~ 1989 The sensitivity of breeding DWVDI EYALL)Es golden plovers Pllt~t~i~~lisapricarin to human intruders Bird Study 3649-55

Ykluc I ExNDDWYt1IIEU 1990 Recreational disturbance of breeding golden plovers Plucic~lis c1Jnicrlrius Biological Consenation 51 243-262

ZIILEK21 H C ZlhN4ND 5IJ ampLUFREDO 1993 Boulder Open Space visitation study Final report prepared for Cir) of Boul- der Open Space-Real Estate Department Boulder Colorado LISA

Scott G Miller received his BS in ivildlite ecology trom the University of Wisconsin-Madison and his MS in ivildlite biol- ogy from Colorado State University His research interests ha~e focused on how landscape ~nodiiications intluence wildliie and vegetative conlmunities and also the impacts of outdoor recre- ation on wildlife Currently Scott coordinates the United States Fish dnd Wildlife Services Partners ior Fish and ildlife Pro-gram in the San Luis Valley Colorado where he restores-enhances wetlands on private property Richard L Knight is a professor in the Department of Fishery and Wildlife Biology at Colorado State University His research interests ~nclude inves- tigating the impacts of outdoor recreation on wildlife and how rural housing developnlents attect Lvildlife communities Rick also devotes much time to preserving ranch land In the LVest Clinton K Miller received his BS in vildlife ecology irom the Universitv of Wisconsin-Madison and his MS in ~vildlite biol- ogy from Colorado State University As the ~vildlite biologist for the city of Boulder Colorado he has dealt heavily with issues of how to manage for both outdoor recreation and vildliie Currently he works for The Nature Conservancy in east- ern South Dakota R Associate editor Bright ampA

You have printed the following article

Wildlife Responses to Pedestrians and DogsScott G Miller Richard L Knight Clinton K MillerWildlife Society Bulletin Vol 29 No 1 (Spring 2001) pp 124-132Stable URL

httplinksjstororgsicisici=0091-76482820012129293A13C1243AWRTPAD3E20CO3B2-0

This article references the following linked citations If you are trying to access articles from anoff-campus location you may be required to first logon via your library web site to access JSTOR Pleasevisit your librarys website or contact a librarian to learn about options for remote access to JSTOR

Literature cited

Variation in Coyote Diets Associated with Season and Successional Changes in VegetationWilliam F Andelt John G Kie Frederick F Knowlton Kean CardwellThe Journal of Wildlife Management Vol 51 No 2 (Apr 1987) pp 273-277Stable URL

httplinksjstororgsicisici=0022-541X2819870429513A23C2733AVICDAW3E20CO3B2-I

Effects of Nonconsumptive Recreation on Wildlife A ReviewStephen A Boyle Fred B SamsonWildlife Society Bulletin Vol 13 No 2 (Summer 1985) pp 110-116Stable URL

httplinksjstororgsicisici=0091-76482819852229133A23C1103AEONROW3E20CO3B2-X

Disturbance Behaviour of Chamois in an Alpine Tourist Area of AustriaJoseph HamrMountain Research and Development Vol 8 No 1 (Feb 1988) pp 65-73Stable URL

httplinksjstororgsicisici=0276-4741281988022983A13C653ADBOCIA3E20CO3B2-P

Human Attitudes and Mountain Sheep in a Wilderness SettingLisa K Harris Paul R Krausman William W ShawWildlife Society Bulletin Vol 23 No 1 (Spring 1995) pp 66-72Stable URL

httplinksjstororgsicisici=0091-76482819952129233A13C663AHAAMSI3E20CO3B2-M

httpwwwjstororg

LINKED CITATIONS- Page 1 of 3 -

1 30 uiildlijk Society Rzrlletin 2 0 0 129(1) 124 - 132

Pedestrian alone (on-tra~l) Dog on leash (on-trail) a pedestrian 1s the addi- 1 00 tive Factor Thls is further

o 80 supported by the fact that the area of influence did

0 60 not differ between the

0 40 0 40 pedestrian-alone and dog- o 20 o 20 on-leash treatments for

0000 20 40 60 80 100 o 00

0 20 40 60 80 100 American robins in forest either on- or

the off--3 Pedestrian-alone (off-trail) Dog on leash (off-trail) trail

For all species in our study area of influence flush distance and dis-tance moved were almost always greater when activities occurred off-trail

0 00 o 00 versus when the same 0 20 40 60 60 100 0 20 40 60 80 100

activih occurred on-trail Recreational use occurred

Pedestrian-alone (on-trail) Dog on leash (on-trail) on our off-trail study sites but was sporadic con-versely recreational use on trails was common (City of Boulder Open Space 1996) Because recreational activities oc- curring on-trail were fre-quent and spatially pre-

Pedestrian alone (off-trail) Dog on leash (off-trail) dictable animals had likely habituated to activi- ty in these locations Off- trail recreation however was infrequent and spa-tially unpredictable Thus animals were not accus-tomed to activity in these areas resulting in the

Meters from trail (for on-trail) or line of movement (for off-trail) greater area of influence

Fiugre 2 Predicted probability of an American robin flushing to treatments in forests during flush distance and dis-1996 City ol Boulder Open Space Boulder Colorado (dashed lines indicate 95 CI) tance moved In Switzer-

land a study of marmots revealed similar results

birds (Leach and Frazier 1953Andelt et al 1987) (Mainini et al 1993) They found that marmots these bird species may not have perceived dogs as exhibited the greatest response to hikers when hik- an important threat Alternatively dogs may pose a ers strayed away from trails Cooke (1 980) Yalden different kind of threat than a pedestrian and birds andYalden (1989) Burger and Gochfeld (1991) and may hold their position until the last moment Kenny and Knight (1992) showed that in areas attempting to remain undetected Because the area where human activity was common and frequent of influence was generally the smallest for the dog- birds were less disturbed than those in areas where alone treatments and because there were no signif- humans were uncommon Likewise Schultz and icant differences between the pedestrian-alone and Bailey (1978) MacArthur et al (1982) and Hamr dog-on-leash treatments it appears that presence of (1988) found that large mammals exhibited the

Pedestrians and dogs Miller et al 131

greatest response when human activity was spatial- Temple 1995) Because off-trailtreatments resulted 1y unpredictable in the greatest area of influence for all wildlife in

Unlike the responses of bird species in our study this study recreational use could be restricted mule deer exhibited the greatest response when a (through education and enforcement) to trails as a dog was present Similar to our results MacArthur way to reduce impacts However because negative et al (1979 1982) and Mainini et al (1993) found impacts occur even from on-trail use number and that lno~lntainsheep and nyarmots respectively spatial arrallgement of trails must be considered in exhibited heightened responses when dogs were conservation planning Furthermore kcause QPe

present ~ l thoughCit)of Boulder Open Space reg- of recreational activity influenced the magnitude of

ulations recluire that dogs be under control wildlife response managers could restrict certain

there were no leash laws 011 our study sites and recreatiollal activities such as prohibiting dogs in

dogs are known to brass alld attack deer (person- some areas or requiring dogs to be leashed Pdrti-

obsemtion) Because dogs can kill deer tiolling the landscape into recreation zones allow-

1953br ick 1969LOWryand Mcampthur 1978) arid ing certain activities in some zones while restrict-

because canicls have on deer tllroLlghollt ing them in others may aid in reducing conflicts

their evolutionary history we assume that deer with sensitive species

have become sensitized to the presence of dogs People are often not aware of h o ~ vtheir activities

explailling the greater reaction when a pedestrian affect wildlife even if the see animals respond to

was accompanied by a dog their actions (Stalmaster and Kaiser 1998) Even F ~ ) ~the species measured ill our studythe area of tllough the dog-alone treatment resulted in the

influence was slnaller whetl treatments occurred smallest area of influence for grassland birds in our

on-trail than off-trail However all species appeared study area of influence will increase if recreation-

to have a threshold of tolerance to disturbance ists alloartheir dogs to roatn amrayfrom a trail Addi-tionally in our study we did not stop and view thcbased on distance with a greater flush response subjects for extended periods of time or attempt to(and alert response for mule deer) when wildlife move toward them Behaviors of this kind are com-

were close to trails An earlier study on the same moll among nature viewers and could lead to ele-

area revealed a positive correlation between abun-vated wildlife responses (mein 1993)

dance of some bird species nest occurrence and Recreationists are more likely to support restric-

nest success with distance from trails r miller et al tions if they understand how wildlife will benefit

1998) The authors felt that this correlation was in (Purcly et al 1987Harris et al 1995) By emphasizingpart a result of recreational actirih and the associ- how human affect wildliie people can ass-ated disturbance Even though the area of influ- ciate their actions or harming encc for all species was sn~alleron-trail versus off- animal populations and begin to develop a conser-a-trail on-trail activities may still constitute an Such an ethic call minilllize important source of disturbance Thus our results wildlife-human conflicts occurring in llatural suggest that human activities may displace wildlife areas ( ~ ~ i ~ h ~and ~~~~l~ 1995) Klein(1993) found and reduce fitness in local wildlife populations A-hat visitors who spoke to wildlife refilge personnel mentioned earlieroff-trailrecreational use was spo- less likely to disturb wildlife than radic However should recreational use away from ho did not llus effectivevisitoreducation can aid trails increase displacelllent of wildlife may ~llti- in developing a collsemation ethic T13rough edllca-mately result Experiments conducted in forested tion land managers can inform recreatiollists how areas of Kyoming without trails Support this con- their activities affectwildlife and how the- can mod-clusion (Gutzwiller et al 1994 Riffell et al 1996 ify their behavior to minimize impacts Gutzwiller et al 1997) showing that recreational activities away from trails resulted in altered behav- Acknoudedgments We thank Madison ancl Bo for

ior and displacement of birds their assistance in the field We are grateful to the City of Boulder Open Space Department for finan-cial support

Management implications Land managers can use spatial and behavioral Literature cited

restrictions in management to L k A l gt ~ I ~ VFJ ( KIFF F K U O U T I O A ~ I )K C ~ I ) Ulili 19X7 Varj-istence of wildlife and recreationists (Knight and ation in coote diets associatecl with season and successional

132 WtXdlt~2Society Bulletin 200123(1) 124- 132

RA GEISSAZID 1982 Cardiac and 2-3-27 behavioral responses of mountain sheep to human distur-

BARICK 1969 Deer predation in North Carolina and other bance Journal of Wildlife Management 46 35 1-358

changes in vegetation Journal of Wildlife Management 51 RL~C~RTHLIII R HJOHYSTOU

F B southeastern states Pages 25-31 it2 White-tailed deer in the ~ZIINISIBI NELIFWL l I ~ ( o I SAUD 11 1993 Behaviour of marmots southern forest habitat Proceeding of a symposi~un United lfnnnota nlarmota under the influence of different hiking States Forest Service Southern Forest Experiment Station activities Biological Consen-ation 64 161-164 Nacogdoches Texas USA MILLER AVLI C K MIILER S G R L K~IGHT 1998 Influence of

BOWERSR R 1953 The free-nmnning dog menace Virginia recreational trails on breeding bird communities Ecological Wildlife 14 5- Applications 8 162-169

BOILE S A AND I B SAISON 1985 Effects of nonconsumptive PLRDIK G G R GOFIDJDLCKERGA P~VEK~UTY ~UL~ NACol-recreation on wildlife a review Wildlife Society Bulletin 13 x~111 198 A guide to managing 11umtn activity 011 Nation-110-116 al Wildlife Refuges Human Dimensions Resources Unit

BURGER M GOCHFELD Department of Natural Resources Cornell Lrniversih- Ithaca JAUI) 1991 Human distance and birds tol- erance and response distances of resident and migrant New York ITSA species in India Environmental Consenration 18 158-165 RIFFELL AUD ANI)ERSO~ DoesS K K J GITZ~TLLER S 1 1996

C m OF B ~ L L D E ROPE^ SFACE 1996 North Boulder Valley inven- repeated human intrusion cause c~~mulative declines in avian tory report City of Boulder Open Space Department Boul- richness and abundance Ecological Applications 6 492-505 der Colorado USA SAS IU5TlTl TE Isc 1988 SASSTAT I-sers Guide Release 603

Coo~cAS 1980 Observations on how close certain passerine Edition SAS Institute Inc Cay North Carolina USA species will tolerate an approaching human in rural and sub- SAS Iha~rrr rh IYC 1993 SAS Technical Report p-243 SASSTAT urban areas Biological Conservation 1885-88 Software the GENRlOD procedure Release 609 SAS Insti-

GLTZWTLLERK J E A KROESES H ~VUERSOU C A WILKINS AND tute Inc Cay North Carolina USA 199 Does human intrusion alter the seasonal timing of avian song during breeding periods Auk 114 55-65

Gri ZWIILER K L CLE~IEUTSKJ RT W I E D E ~ ~ N N ANI) 5 HAuDIR- sos 1994 Effects of human intrusions on song occurrence and singing consistency in subalpine birds Auk 11 1 28-37

H ~ M K 1988 Disturbance behaviour of chamois in an alpine J tourist area of Austria Mountain Research and Development 865-73

HARRIS AUD WW SHAY L K R R K R ~ L ~ S ~ M N 1995 Human atti- tudes and mountain sheep in a wilderness setting Wildlife Society Bulletin 2366-2

KEXWSA R L KNIGHT 1992 Flight distances of black- billed magpies in different regimes of human densih and persecution Condor 94 545- 547

KLEIN 1993 Waterbird behavioral responses to human dis- M L turbances Wildlife Society Bulletin 21 31 -39

KSIGHTR LAYD D N COIE 1995 Factors that influence wildlife responses to recreationists Pages 1-9 in Wildlife and recreationists coexistence through research and manage- ment R L Knight and K J Gutnl-iller editors Island Cov- elo California USA

KNIGHT 1995 Wildlife and recreation- R L~ N I )K J GL TZWILLER

ists coexistence through research and management R L Knight and K Gutzwiller editors Island Covelo California USA

KYIGHT 1995 Wildlife and recreationists R L A ~ L )SATEMPLE coexistence through management Pages 32-333 in Wildlife and recreationists coexistence through research and management R L Knight and K J Gutn7iller editors Island Corelo California USA

LWCHH R IUD W H FMIER 1953 A study of the possible extent of predation on heavy concentrations of valley quail with special reference to the bobcat California Fish and Game 39 52-538

LOWRY 1978 Domestic dogs as pred- DAAND K L ampI(-ARTHUR ators on deer Wildlife Society Bulletin 6 38-39

RIACARTHLR GEIST Factors influ- RA R H JOHNSTOSA~DV 19-9 encing heart rate in free-ranging bighorn sheep a physiolog-ical approach to the study of wildlife harassment Canadian Journal of Zoolog)- 572010-2021

SCHLLTZR DID JA BAILII 198 R~S~OIISCSof national park elk to human activity Journal of Wildlife lanagenlent i2 91-100

STampIJI~STFR J L KAISER Effect of recreational Y1 AD 1998 activity on wintering bald eagles Wiltllife Monograph 13-

WH~-laakRD ampnu RL KUIGHT 1998 Understanding wildlife responses to humans Wildlife Society Bulletin 26 312-31

YALDE~ 1989 The sensitivity of breeding DWVDI EYALL)Es golden plovers Pllt~t~i~~lisapricarin to human intruders Bird Study 3649-55

Ykluc I ExNDDWYt1IIEU 1990 Recreational disturbance of breeding golden plovers Plucic~lis c1Jnicrlrius Biological Consenation 51 243-262

ZIILEK21 H C ZlhN4ND 5IJ ampLUFREDO 1993 Boulder Open Space visitation study Final report prepared for Cir) of Boul- der Open Space-Real Estate Department Boulder Colorado LISA

Scott G Miller received his BS in ivildlite ecology trom the University of Wisconsin-Madison and his MS in ivildlite biol- ogy from Colorado State University His research interests ha~e focused on how landscape ~nodiiications intluence wildliie and vegetative conlmunities and also the impacts of outdoor recre- ation on wildlife Currently Scott coordinates the United States Fish dnd Wildlife Services Partners ior Fish and ildlife Pro-gram in the San Luis Valley Colorado where he restores-enhances wetlands on private property Richard L Knight is a professor in the Department of Fishery and Wildlife Biology at Colorado State University His research interests ~nclude inves- tigating the impacts of outdoor recreation on wildlife and how rural housing developnlents attect Lvildlife communities Rick also devotes much time to preserving ranch land In the LVest Clinton K Miller received his BS in vildlife ecology irom the Universitv of Wisconsin-Madison and his MS in ~vildlite biol- ogy from Colorado State University As the ~vildlite biologist for the city of Boulder Colorado he has dealt heavily with issues of how to manage for both outdoor recreation and vildliie Currently he works for The Nature Conservancy in east- ern South Dakota R Associate editor Bright ampA

You have printed the following article

Wildlife Responses to Pedestrians and DogsScott G Miller Richard L Knight Clinton K MillerWildlife Society Bulletin Vol 29 No 1 (Spring 2001) pp 124-132Stable URL

httplinksjstororgsicisici=0091-76482820012129293A13C1243AWRTPAD3E20CO3B2-0

This article references the following linked citations If you are trying to access articles from anoff-campus location you may be required to first logon via your library web site to access JSTOR Pleasevisit your librarys website or contact a librarian to learn about options for remote access to JSTOR

Literature cited

Variation in Coyote Diets Associated with Season and Successional Changes in VegetationWilliam F Andelt John G Kie Frederick F Knowlton Kean CardwellThe Journal of Wildlife Management Vol 51 No 2 (Apr 1987) pp 273-277Stable URL

httplinksjstororgsicisici=0022-541X2819870429513A23C2733AVICDAW3E20CO3B2-I

Effects of Nonconsumptive Recreation on Wildlife A ReviewStephen A Boyle Fred B SamsonWildlife Society Bulletin Vol 13 No 2 (Summer 1985) pp 110-116Stable URL

httplinksjstororgsicisici=0091-76482819852229133A23C1103AEONROW3E20CO3B2-X

Disturbance Behaviour of Chamois in an Alpine Tourist Area of AustriaJoseph HamrMountain Research and Development Vol 8 No 1 (Feb 1988) pp 65-73Stable URL

httplinksjstororgsicisici=0276-4741281988022983A13C653ADBOCIA3E20CO3B2-P

Human Attitudes and Mountain Sheep in a Wilderness SettingLisa K Harris Paul R Krausman William W ShawWildlife Society Bulletin Vol 23 No 1 (Spring 1995) pp 66-72Stable URL

httplinksjstororgsicisici=0091-76482819952129233A13C663AHAAMSI3E20CO3B2-M

httpwwwjstororg

LINKED CITATIONS- Page 1 of 3 -

Pedestrians and dogs Miller et al 131

greatest response when human activity was spatial- Temple 1995) Because off-trailtreatments resulted 1y unpredictable in the greatest area of influence for all wildlife in

Unlike the responses of bird species in our study this study recreational use could be restricted mule deer exhibited the greatest response when a (through education and enforcement) to trails as a dog was present Similar to our results MacArthur way to reduce impacts However because negative et al (1979 1982) and Mainini et al (1993) found impacts occur even from on-trail use number and that lno~lntainsheep and nyarmots respectively spatial arrallgement of trails must be considered in exhibited heightened responses when dogs were conservation planning Furthermore kcause QPe

present ~ l thoughCit)of Boulder Open Space reg- of recreational activity influenced the magnitude of

ulations recluire that dogs be under control wildlife response managers could restrict certain

there were no leash laws 011 our study sites and recreatiollal activities such as prohibiting dogs in

dogs are known to brass alld attack deer (person- some areas or requiring dogs to be leashed Pdrti-

obsemtion) Because dogs can kill deer tiolling the landscape into recreation zones allow-

1953br ick 1969LOWryand Mcampthur 1978) arid ing certain activities in some zones while restrict-

because canicls have on deer tllroLlghollt ing them in others may aid in reducing conflicts

their evolutionary history we assume that deer with sensitive species

have become sensitized to the presence of dogs People are often not aware of h o ~ vtheir activities

explailling the greater reaction when a pedestrian affect wildlife even if the see animals respond to

was accompanied by a dog their actions (Stalmaster and Kaiser 1998) Even F ~ ) ~the species measured ill our studythe area of tllough the dog-alone treatment resulted in the

influence was slnaller whetl treatments occurred smallest area of influence for grassland birds in our

on-trail than off-trail However all species appeared study area of influence will increase if recreation-

to have a threshold of tolerance to disturbance ists alloartheir dogs to roatn amrayfrom a trail Addi-tionally in our study we did not stop and view thcbased on distance with a greater flush response subjects for extended periods of time or attempt to(and alert response for mule deer) when wildlife move toward them Behaviors of this kind are com-

were close to trails An earlier study on the same moll among nature viewers and could lead to ele-

area revealed a positive correlation between abun-vated wildlife responses (mein 1993)

dance of some bird species nest occurrence and Recreationists are more likely to support restric-

nest success with distance from trails r miller et al tions if they understand how wildlife will benefit

1998) The authors felt that this correlation was in (Purcly et al 1987Harris et al 1995) By emphasizingpart a result of recreational actirih and the associ- how human affect wildliie people can ass-ated disturbance Even though the area of influ- ciate their actions or harming encc for all species was sn~alleron-trail versus off- animal populations and begin to develop a conser-a-trail on-trail activities may still constitute an Such an ethic call minilllize important source of disturbance Thus our results wildlife-human conflicts occurring in llatural suggest that human activities may displace wildlife areas ( ~ ~ i ~ h ~and ~~~~l~ 1995) Klein(1993) found and reduce fitness in local wildlife populations A-hat visitors who spoke to wildlife refilge personnel mentioned earlieroff-trailrecreational use was spo- less likely to disturb wildlife than radic However should recreational use away from ho did not llus effectivevisitoreducation can aid trails increase displacelllent of wildlife may ~llti- in developing a collsemation ethic T13rough edllca-mately result Experiments conducted in forested tion land managers can inform recreatiollists how areas of Kyoming without trails Support this con- their activities affectwildlife and how the- can mod-clusion (Gutzwiller et al 1994 Riffell et al 1996 ify their behavior to minimize impacts Gutzwiller et al 1997) showing that recreational activities away from trails resulted in altered behav- Acknoudedgments We thank Madison ancl Bo for

ior and displacement of birds their assistance in the field We are grateful to the City of Boulder Open Space Department for finan-cial support

Management implications Land managers can use spatial and behavioral Literature cited

restrictions in management to L k A l gt ~ I ~ VFJ ( KIFF F K U O U T I O A ~ I )K C ~ I ) Ulili 19X7 Varj-istence of wildlife and recreationists (Knight and ation in coote diets associatecl with season and successional

132 WtXdlt~2Society Bulletin 200123(1) 124- 132

RA GEISSAZID 1982 Cardiac and 2-3-27 behavioral responses of mountain sheep to human distur-

BARICK 1969 Deer predation in North Carolina and other bance Journal of Wildlife Management 46 35 1-358

changes in vegetation Journal of Wildlife Management 51 RL~C~RTHLIII R HJOHYSTOU

F B southeastern states Pages 25-31 it2 White-tailed deer in the ~ZIINISIBI NELIFWL l I ~ ( o I SAUD 11 1993 Behaviour of marmots southern forest habitat Proceeding of a symposi~un United lfnnnota nlarmota under the influence of different hiking States Forest Service Southern Forest Experiment Station activities Biological Consen-ation 64 161-164 Nacogdoches Texas USA MILLER AVLI C K MIILER S G R L K~IGHT 1998 Influence of

BOWERSR R 1953 The free-nmnning dog menace Virginia recreational trails on breeding bird communities Ecological Wildlife 14 5- Applications 8 162-169

BOILE S A AND I B SAISON 1985 Effects of nonconsumptive PLRDIK G G R GOFIDJDLCKERGA P~VEK~UTY ~UL~ NACol-recreation on wildlife a review Wildlife Society Bulletin 13 x~111 198 A guide to managing 11umtn activity 011 Nation-110-116 al Wildlife Refuges Human Dimensions Resources Unit

BURGER M GOCHFELD Department of Natural Resources Cornell Lrniversih- Ithaca JAUI) 1991 Human distance and birds tol- erance and response distances of resident and migrant New York ITSA species in India Environmental Consenration 18 158-165 RIFFELL AUD ANI)ERSO~ DoesS K K J GITZ~TLLER S 1 1996

C m OF B ~ L L D E ROPE^ SFACE 1996 North Boulder Valley inven- repeated human intrusion cause c~~mulative declines in avian tory report City of Boulder Open Space Department Boul- richness and abundance Ecological Applications 6 492-505 der Colorado USA SAS IU5TlTl TE Isc 1988 SASSTAT I-sers Guide Release 603

Coo~cAS 1980 Observations on how close certain passerine Edition SAS Institute Inc Cay North Carolina USA species will tolerate an approaching human in rural and sub- SAS Iha~rrr rh IYC 1993 SAS Technical Report p-243 SASSTAT urban areas Biological Conservation 1885-88 Software the GENRlOD procedure Release 609 SAS Insti-

GLTZWTLLERK J E A KROESES H ~VUERSOU C A WILKINS AND tute Inc Cay North Carolina USA 199 Does human intrusion alter the seasonal timing of avian song during breeding periods Auk 114 55-65

Gri ZWIILER K L CLE~IEUTSKJ RT W I E D E ~ ~ N N ANI) 5 HAuDIR- sos 1994 Effects of human intrusions on song occurrence and singing consistency in subalpine birds Auk 11 1 28-37

H ~ M K 1988 Disturbance behaviour of chamois in an alpine J tourist area of Austria Mountain Research and Development 865-73

HARRIS AUD WW SHAY L K R R K R ~ L ~ S ~ M N 1995 Human atti- tudes and mountain sheep in a wilderness setting Wildlife Society Bulletin 2366-2

KEXWSA R L KNIGHT 1992 Flight distances of black- billed magpies in different regimes of human densih and persecution Condor 94 545- 547

KLEIN 1993 Waterbird behavioral responses to human dis- M L turbances Wildlife Society Bulletin 21 31 -39

KSIGHTR LAYD D N COIE 1995 Factors that influence wildlife responses to recreationists Pages 1-9 in Wildlife and recreationists coexistence through research and manage- ment R L Knight and K J Gutnl-iller editors Island Cov- elo California USA

KNIGHT 1995 Wildlife and recreation- R L~ N I )K J GL TZWILLER

ists coexistence through research and management R L Knight and K Gutzwiller editors Island Covelo California USA

KYIGHT 1995 Wildlife and recreationists R L A ~ L )SATEMPLE coexistence through management Pages 32-333 in Wildlife and recreationists coexistence through research and management R L Knight and K J Gutn7iller editors Island Corelo California USA

LWCHH R IUD W H FMIER 1953 A study of the possible extent of predation on heavy concentrations of valley quail with special reference to the bobcat California Fish and Game 39 52-538

LOWRY 1978 Domestic dogs as pred- DAAND K L ampI(-ARTHUR ators on deer Wildlife Society Bulletin 6 38-39

RIACARTHLR GEIST Factors influ- RA R H JOHNSTOSA~DV 19-9 encing heart rate in free-ranging bighorn sheep a physiolog-ical approach to the study of wildlife harassment Canadian Journal of Zoolog)- 572010-2021

SCHLLTZR DID JA BAILII 198 R~S~OIISCSof national park elk to human activity Journal of Wildlife lanagenlent i2 91-100

STampIJI~STFR J L KAISER Effect of recreational Y1 AD 1998 activity on wintering bald eagles Wiltllife Monograph 13-

WH~-laakRD ampnu RL KUIGHT 1998 Understanding wildlife responses to humans Wildlife Society Bulletin 26 312-31

YALDE~ 1989 The sensitivity of breeding DWVDI EYALL)Es golden plovers Pllt~t~i~~lisapricarin to human intruders Bird Study 3649-55

Ykluc I ExNDDWYt1IIEU 1990 Recreational disturbance of breeding golden plovers Plucic~lis c1Jnicrlrius Biological Consenation 51 243-262

ZIILEK21 H C ZlhN4ND 5IJ ampLUFREDO 1993 Boulder Open Space visitation study Final report prepared for Cir) of Boul- der Open Space-Real Estate Department Boulder Colorado LISA

Scott G Miller received his BS in ivildlite ecology trom the University of Wisconsin-Madison and his MS in ivildlite biol- ogy from Colorado State University His research interests ha~e focused on how landscape ~nodiiications intluence wildliie and vegetative conlmunities and also the impacts of outdoor recre- ation on wildlife Currently Scott coordinates the United States Fish dnd Wildlife Services Partners ior Fish and ildlife Pro-gram in the San Luis Valley Colorado where he restores-enhances wetlands on private property Richard L Knight is a professor in the Department of Fishery and Wildlife Biology at Colorado State University His research interests ~nclude inves- tigating the impacts of outdoor recreation on wildlife and how rural housing developnlents attect Lvildlife communities Rick also devotes much time to preserving ranch land In the LVest Clinton K Miller received his BS in vildlife ecology irom the Universitv of Wisconsin-Madison and his MS in ~vildlite biol- ogy from Colorado State University As the ~vildlite biologist for the city of Boulder Colorado he has dealt heavily with issues of how to manage for both outdoor recreation and vildliie Currently he works for The Nature Conservancy in east- ern South Dakota R Associate editor Bright ampA

You have printed the following article

Wildlife Responses to Pedestrians and DogsScott G Miller Richard L Knight Clinton K MillerWildlife Society Bulletin Vol 29 No 1 (Spring 2001) pp 124-132Stable URL

httplinksjstororgsicisici=0091-76482820012129293A13C1243AWRTPAD3E20CO3B2-0

This article references the following linked citations If you are trying to access articles from anoff-campus location you may be required to first logon via your library web site to access JSTOR Pleasevisit your librarys website or contact a librarian to learn about options for remote access to JSTOR

Literature cited

Variation in Coyote Diets Associated with Season and Successional Changes in VegetationWilliam F Andelt John G Kie Frederick F Knowlton Kean CardwellThe Journal of Wildlife Management Vol 51 No 2 (Apr 1987) pp 273-277Stable URL

httplinksjstororgsicisici=0022-541X2819870429513A23C2733AVICDAW3E20CO3B2-I

Effects of Nonconsumptive Recreation on Wildlife A ReviewStephen A Boyle Fred B SamsonWildlife Society Bulletin Vol 13 No 2 (Summer 1985) pp 110-116Stable URL

httplinksjstororgsicisici=0091-76482819852229133A23C1103AEONROW3E20CO3B2-X

Disturbance Behaviour of Chamois in an Alpine Tourist Area of AustriaJoseph HamrMountain Research and Development Vol 8 No 1 (Feb 1988) pp 65-73Stable URL

httplinksjstororgsicisici=0276-4741281988022983A13C653ADBOCIA3E20CO3B2-P

Human Attitudes and Mountain Sheep in a Wilderness SettingLisa K Harris Paul R Krausman William W ShawWildlife Society Bulletin Vol 23 No 1 (Spring 1995) pp 66-72Stable URL

httplinksjstororgsicisici=0091-76482819952129233A13C663AHAAMSI3E20CO3B2-M

httpwwwjstororg

LINKED CITATIONS- Page 1 of 3 -

132 WtXdlt~2Society Bulletin 200123(1) 124- 132

RA GEISSAZID 1982 Cardiac and 2-3-27 behavioral responses of mountain sheep to human distur-

BARICK 1969 Deer predation in North Carolina and other bance Journal of Wildlife Management 46 35 1-358

changes in vegetation Journal of Wildlife Management 51 RL~C~RTHLIII R HJOHYSTOU

F B southeastern states Pages 25-31 it2 White-tailed deer in the ~ZIINISIBI NELIFWL l I ~ ( o I SAUD 11 1993 Behaviour of marmots southern forest habitat Proceeding of a symposi~un United lfnnnota nlarmota under the influence of different hiking States Forest Service Southern Forest Experiment Station activities Biological Consen-ation 64 161-164 Nacogdoches Texas USA MILLER AVLI C K MIILER S G R L K~IGHT 1998 Influence of

BOWERSR R 1953 The free-nmnning dog menace Virginia recreational trails on breeding bird communities Ecological Wildlife 14 5- Applications 8 162-169

BOILE S A AND I B SAISON 1985 Effects of nonconsumptive PLRDIK G G R GOFIDJDLCKERGA P~VEK~UTY ~UL~ NACol-recreation on wildlife a review Wildlife Society Bulletin 13 x~111 198 A guide to managing 11umtn activity 011 Nation-110-116 al Wildlife Refuges Human Dimensions Resources Unit

BURGER M GOCHFELD Department of Natural Resources Cornell Lrniversih- Ithaca JAUI) 1991 Human distance and birds tol- erance and response distances of resident and migrant New York ITSA species in India Environmental Consenration 18 158-165 RIFFELL AUD ANI)ERSO~ DoesS K K J GITZ~TLLER S 1 1996

C m OF B ~ L L D E ROPE^ SFACE 1996 North Boulder Valley inven- repeated human intrusion cause c~~mulative declines in avian tory report City of Boulder Open Space Department Boul- richness and abundance Ecological Applications 6 492-505 der Colorado USA SAS IU5TlTl TE Isc 1988 SASSTAT I-sers Guide Release 603

Coo~cAS 1980 Observations on how close certain passerine Edition SAS Institute Inc Cay North Carolina USA species will tolerate an approaching human in rural and sub- SAS Iha~rrr rh IYC 1993 SAS Technical Report p-243 SASSTAT urban areas Biological Conservation 1885-88 Software the GENRlOD procedure Release 609 SAS Insti-

GLTZWTLLERK J E A KROESES H ~VUERSOU C A WILKINS AND tute Inc Cay North Carolina USA 199 Does human intrusion alter the seasonal timing of avian song during breeding periods Auk 114 55-65

Gri ZWIILER K L CLE~IEUTSKJ RT W I E D E ~ ~ N N ANI) 5 HAuDIR- sos 1994 Effects of human intrusions on song occurrence and singing consistency in subalpine birds Auk 11 1 28-37

H ~ M K 1988 Disturbance behaviour of chamois in an alpine J tourist area of Austria Mountain Research and Development 865-73

HARRIS AUD WW SHAY L K R R K R ~ L ~ S ~ M N 1995 Human atti- tudes and mountain sheep in a wilderness setting Wildlife Society Bulletin 2366-2

KEXWSA R L KNIGHT 1992 Flight distances of black- billed magpies in different regimes of human densih and persecution Condor 94 545- 547

KLEIN 1993 Waterbird behavioral responses to human dis- M L turbances Wildlife Society Bulletin 21 31 -39

KSIGHTR LAYD D N COIE 1995 Factors that influence wildlife responses to recreationists Pages 1-9 in Wildlife and recreationists coexistence through research and manage- ment R L Knight and K J Gutnl-iller editors Island Cov- elo California USA

KNIGHT 1995 Wildlife and recreation- R L~ N I )K J GL TZWILLER

ists coexistence through research and management R L Knight and K Gutzwiller editors Island Covelo California USA

KYIGHT 1995 Wildlife and recreationists R L A ~ L )SATEMPLE coexistence through management Pages 32-333 in Wildlife and recreationists coexistence through research and management R L Knight and K J Gutn7iller editors Island Corelo California USA

LWCHH R IUD W H FMIER 1953 A study of the possible extent of predation on heavy concentrations of valley quail with special reference to the bobcat California Fish and Game 39 52-538

LOWRY 1978 Domestic dogs as pred- DAAND K L ampI(-ARTHUR ators on deer Wildlife Society Bulletin 6 38-39

RIACARTHLR GEIST Factors influ- RA R H JOHNSTOSA~DV 19-9 encing heart rate in free-ranging bighorn sheep a physiolog-ical approach to the study of wildlife harassment Canadian Journal of Zoolog)- 572010-2021

SCHLLTZR DID JA BAILII 198 R~S~OIISCSof national park elk to human activity Journal of Wildlife lanagenlent i2 91-100

STampIJI~STFR J L KAISER Effect of recreational Y1 AD 1998 activity on wintering bald eagles Wiltllife Monograph 13-

WH~-laakRD ampnu RL KUIGHT 1998 Understanding wildlife responses to humans Wildlife Society Bulletin 26 312-31

YALDE~ 1989 The sensitivity of breeding DWVDI EYALL)Es golden plovers Pllt~t~i~~lisapricarin to human intruders Bird Study 3649-55

Ykluc I ExNDDWYt1IIEU 1990 Recreational disturbance of breeding golden plovers Plucic~lis c1Jnicrlrius Biological Consenation 51 243-262

ZIILEK21 H C ZlhN4ND 5IJ ampLUFREDO 1993 Boulder Open Space visitation study Final report prepared for Cir) of Boul- der Open Space-Real Estate Department Boulder Colorado LISA

Scott G Miller received his BS in ivildlite ecology trom the University of Wisconsin-Madison and his MS in ivildlite biol- ogy from Colorado State University His research interests ha~e focused on how landscape ~nodiiications intluence wildliie and vegetative conlmunities and also the impacts of outdoor recre- ation on wildlife Currently Scott coordinates the United States Fish dnd Wildlife Services Partners ior Fish and ildlife Pro-gram in the San Luis Valley Colorado where he restores-enhances wetlands on private property Richard L Knight is a professor in the Department of Fishery and Wildlife Biology at Colorado State University His research interests ~nclude inves- tigating the impacts of outdoor recreation on wildlife and how rural housing developnlents attect Lvildlife communities Rick also devotes much time to preserving ranch land In the LVest Clinton K Miller received his BS in vildlife ecology irom the Universitv of Wisconsin-Madison and his MS in ~vildlite biol- ogy from Colorado State University As the ~vildlite biologist for the city of Boulder Colorado he has dealt heavily with issues of how to manage for both outdoor recreation and vildliie Currently he works for The Nature Conservancy in east- ern South Dakota R Associate editor Bright ampA

You have printed the following article

Wildlife Responses to Pedestrians and DogsScott G Miller Richard L Knight Clinton K MillerWildlife Society Bulletin Vol 29 No 1 (Spring 2001) pp 124-132Stable URL

httplinksjstororgsicisici=0091-76482820012129293A13C1243AWRTPAD3E20CO3B2-0

This article references the following linked citations If you are trying to access articles from anoff-campus location you may be required to first logon via your library web site to access JSTOR Pleasevisit your librarys website or contact a librarian to learn about options for remote access to JSTOR

Literature cited

Variation in Coyote Diets Associated with Season and Successional Changes in VegetationWilliam F Andelt John G Kie Frederick F Knowlton Kean CardwellThe Journal of Wildlife Management Vol 51 No 2 (Apr 1987) pp 273-277Stable URL

httplinksjstororgsicisici=0022-541X2819870429513A23C2733AVICDAW3E20CO3B2-I

Effects of Nonconsumptive Recreation on Wildlife A ReviewStephen A Boyle Fred B SamsonWildlife Society Bulletin Vol 13 No 2 (Summer 1985) pp 110-116Stable URL

httplinksjstororgsicisici=0091-76482819852229133A23C1103AEONROW3E20CO3B2-X

Disturbance Behaviour of Chamois in an Alpine Tourist Area of AustriaJoseph HamrMountain Research and Development Vol 8 No 1 (Feb 1988) pp 65-73Stable URL

httplinksjstororgsicisici=0276-4741281988022983A13C653ADBOCIA3E20CO3B2-P

Human Attitudes and Mountain Sheep in a Wilderness SettingLisa K Harris Paul R Krausman William W ShawWildlife Society Bulletin Vol 23 No 1 (Spring 1995) pp 66-72Stable URL

httplinksjstororgsicisici=0091-76482819952129233A13C663AHAAMSI3E20CO3B2-M

httpwwwjstororg

LINKED CITATIONS- Page 1 of 3 -

You have printed the following article

Wildlife Responses to Pedestrians and DogsScott G Miller Richard L Knight Clinton K MillerWildlife Society Bulletin Vol 29 No 1 (Spring 2001) pp 124-132Stable URL

httplinksjstororgsicisici=0091-76482820012129293A13C1243AWRTPAD3E20CO3B2-0

This article references the following linked citations If you are trying to access articles from anoff-campus location you may be required to first logon via your library web site to access JSTOR Pleasevisit your librarys website or contact a librarian to learn about options for remote access to JSTOR

Literature cited

Variation in Coyote Diets Associated with Season and Successional Changes in VegetationWilliam F Andelt John G Kie Frederick F Knowlton Kean CardwellThe Journal of Wildlife Management Vol 51 No 2 (Apr 1987) pp 273-277Stable URL

httplinksjstororgsicisici=0022-541X2819870429513A23C2733AVICDAW3E20CO3B2-I

Effects of Nonconsumptive Recreation on Wildlife A ReviewStephen A Boyle Fred B SamsonWildlife Society Bulletin Vol 13 No 2 (Summer 1985) pp 110-116Stable URL

httplinksjstororgsicisici=0091-76482819852229133A23C1103AEONROW3E20CO3B2-X

Disturbance Behaviour of Chamois in an Alpine Tourist Area of AustriaJoseph HamrMountain Research and Development Vol 8 No 1 (Feb 1988) pp 65-73Stable URL

httplinksjstororgsicisici=0276-4741281988022983A13C653ADBOCIA3E20CO3B2-P

Human Attitudes and Mountain Sheep in a Wilderness SettingLisa K Harris Paul R Krausman William W ShawWildlife Society Bulletin Vol 23 No 1 (Spring 1995) pp 66-72Stable URL

httplinksjstororgsicisici=0091-76482819952129233A13C663AHAAMSI3E20CO3B2-M

httpwwwjstororg

LINKED CITATIONS- Page 1 of 3 -

Waterbird Behavioral Responses to Human DisturbancesMary L KleinWildlife Society Bulletin Vol 21 No 1 (Spring 1993) pp 31-39Stable URL

httplinksjstororgsicisici=0091-76482819932129213A13C313AWBRTHD3E20CO3B2-E

Domestic Dogs as Predators on DeerDwain A Lowry Katherine L McArthurWildlife Society Bulletin Vol 6 No 1 (Spring 1978) pp 38-39Stable URL

httplinksjstororgsicisici=0091-7648281978212963A13C383ADDAPOD3E20CO3B2-Y

Cardiac and Behavioral Responses of Mountain Sheep to Human DisturbanceRobert A MacArthur Valerius Geist Ronald H JohnstonThe Journal of Wildlife Management Vol 46 No 2 (Apr 1982) pp 351-358Stable URL

httplinksjstororgsicisici=0022-541X2819820429463A23C3513ACABROM3E20CO3B2-J

Influence of Recreational Trails on Breeding Bird CommunitiesScott G Miller Richard L Knight Clinton K MillerEcological Applications Vol 8 No 1 (Feb 1998) pp 162-169Stable URL

httplinksjstororgsicisici=1051-0761281998022983A13C1623AIORTOB3E20CO3B2-C

Does Repeated Human Intrusion Cause Cumulative Declines in Avian Richness andAbundanceSamuel K Riffell Kevin J Gutzwiller Stanley H AndersonEcological Applications Vol 6 No 2 (May 1996) pp 492-505Stable URL

httplinksjstororgsicisici=1051-0761281996052963A23C4923ADRHICC3E20CO3B2-B

Responses of National Park Elk to Human ActivityRichard D Schultz James A BaileyThe Journal of Wildlife Management Vol 42 No 1 (Jan 1978) pp 91-100Stable URL

httplinksjstororgsicisici=0022-541X2819780129423A13C913ARONPET3E20CO3B2-2

httpwwwjstororg

LINKED CITATIONS- Page 2 of 3 -

Understanding Wildlife Responses to HumansDoug Whittaker Richard L KnightWildlife Society Bulletin Vol 26 No 2 (Summer 1998) pp 312-317Stable URL

httplinksjstororgsicisici=0091-76482819982229263A23C3123AUWRTH3E20CO3B2-8

httpwwwjstororg

LINKED CITATIONS- Page 3 of 3 -