FGM Samples collected 2 to 4 times per week. Questionnaire based: Number of hiding spots, closest possible visitor proximity, number of enclosure sides visitors can view

Negative impact of number of hiding spots, closest possible proxmity of visitors had a positive relationship with FGMs.

Woodland Park Zoo

Pacing, aggression, social, non visible behaviour

Visitor noise categorised as quite low, moderate, high and extreme. Density recorded on the same scale (groups of numbers)" 8hrs per week for 29 weeks (230hr). Continous focal animal sampling over a 30 minute period. Number and noise of visitors recorded evey one minute.

Number and noise

11m wide x 8m deep x 4m high

Negative effect of visitor number on time spent non visible (highest at lowest visitor density. Male showed increased aggression at higher densities. Female pacing affected by intensity (highest at low levels of talking). Male showed changes in social behaviour in relation to aggression but not clear direction of the effect. Time spent non visible highest at lowest intensity level. No impact on aggression.

Ailuropoda melanoleuca

Exploration, vigilance, locomotion, scent marking, feeding, solitary play, maintenance, manipulation, interaction with humans, stereotypies, social interaction, inactivity, not visible, enclosure use

34 one hour sessions, behaviour recorded at 2minute intervals. Studied one animal at a time

Presence/absence

One indoor and one outdoor enclosure. Indoor 46m2 each . Outdoor enclosures were naturalistic 495m2 each

Percent occurrence of behaviour. SPI for space use

More exploration, feeding, manipulation, stationary and not visible when visitors present. In presence of visitors used back space more frequently than front. Used more of the enclosure when visitors present.

Ursus arctos arctos

Individually housed in semi naturalised enclosures. Female 150m2, male 230m2

More stereotypies, stationary. locomotion and vigialnce when visitors present. In presence of visitors used back space more frequently than front. Used more of the enclosure when visitors present.

Bosque dos Jequitibás zoo

Locomotion, inactivity, basic physiological needs, agonistic threat, exploratory behaviour, vocalisation, directed gaze, in refuge area (OOS)

83hrs of observations captured in two one-week periods. 6hrs per day. Visitor noise and number recorded every 2 minutes.

Number and noise

Effect of noise level and visitor number on behaviour

Behaviour affected but only in the morning. For both individuals noise level impacted on inactivity (higher during higher noise level). Refuge area use was greatest for one indivual during low visitors. Interactions between visitor numbers and noise. Inactivity in one cat higher when high numbers of visitors producing normal or loud noise. For second puma inactivity and directed gaze high when number of visitors high and noise loud.

Fota Wildlife Park

Active (locomotion) or inactive (sleeping/resting) plus social for cubs. Reaction to visitors. Position in enclosure recorded

Data collected during June/July (busy time periods). Observations undertaken in two bouts of 3hrs each day. 10 minute periods with 30 second scans.

Number and noise

Outdoor enclosure

Impact of visitors on behaviour and enclosure use

No impact of visitors on behaviour and enclosure use.

Ailuropoda melanoleuca

Behaviour & Physiology

Behaviour: locomotion, vocalising, door scratching. Physiology: urinary cortisol

Observations for 2hrs 5 afternoons per week. 1 minute samples. Analysed locomotion, honking and door scratching. Urine samples collected each morning. Sound collected every minute and then averaged.

Days characterized by louder levels of noise were associated with increased locomotion, restless manipulation of the exit door of the enclosure, increased scratching and vocalizations indicative of agitation, and/or increased glucocorticoids excreted in urine.

Zoos South Australia

3 months - one tour day and one non-tour day per week

Behaviour (inactive, active species-typical, feeding, conspecific interaction, human interaction, pacing, aggression, out of sight) and proximity to humans

30sec scan samples to look at behaviour before, during and after the tour. Lion and tiger protected contact tour. Cheetah hands on tour. Lion obs: 1hr before, 15 min during, 1hr after. Tiger obs: 30 mins before, 15 mins during, 1hr after. Cheetah: 1hr before, 1hr during, 1hr after

Interaction programme

255m2 outdoor encosure

Inactive lower during tours than pre and post, higher post tour than pre tour. Feeding higher during tours compared to pre and post, higher post tour than pre tour. Pacing higher during tour than pre and post tour, higher pre tour. No change in sp typical behaviour.

four outdoor enclosures 275m2

4500m2 naturalistic enclosure

Inactive highest post tour compared to pre-tour and during tour. Active sp typical behaviour higher pre-tour than post-tour, higher during tour than post-tour. Feeding higher during tours than pre and post tour, higher post tour than pre-tour. HAI higher during tours than pre and post tour. Pacing higher pre tour than during and post. Higher post tour than during. OOS higher during tour than pre tour. More inactive behaviour on tour days vs non tour days.

Parque Zoobotánico and Zoológico Municipal

Behaviour coded as inactivity (sleeping, dozing, attentive, grooming) medium activity (alert, marking, locomotion) and high actvity (playing, sexual behaviour, stereotypies, agonistic behaviour)

1 minute recording every 5 minutes. one scan per recording. 150 scans. 15hrs of activity per animal

Open/closed and number

During zoo open mornings less active spending more time sleeping; they played and walked less . agonistic behaviour only when visitors were present . No impact of visitor density.

Greater activity - spent less time dozing and more time grooming and walking in the presence of visitors. more active on zoo open days and even directed some playful behaviour towards visitors No impact of visitor density. On zoo open days the jaguar made more use of areas closer to visitors

During zoo open mornings less active spending more time sleeping; they played and walked less . No impact of visitor density. On zoo open days spent more time in retreat spaces and hiding places and less time in areas closer to the public

leopardus pardalis,

During zoo open mornings less active spending more time sleeping; they played and walked less and devoted less time to abnormal stereotypic behaviours. agonistic behaviour only when visitors were present .No impact of visitor density. On zoo open days spent more time in retreat spaces and hiding places and less time in areas closer to the public

panthero leo persica

Less abnormal stereotypic behaviours during zoo open mornings. agonistic behaviour only when visitors were present .No impact of visitor density.

10hrs (24 days over 6 weeks)

Swimming under water, swimming with head above water, visually scanning environment, resting partially on land, social interactions

2 to 3 10min observations per day. Total 61 observations

16m2 pool with beach

compared 7 highest days of visitors with 7 lowest days of visitors and compared to seal behaviour

On busy days: more time spent swimming under water, less time scanning the environment and resting on land. No difference in swimming head above water or social behaviour.

2M 2F at NEX, 2M 1F at BZ

Brasilia Zoo (BZ) & “No Extinction” (NEX)

2 months, 9 samples per jaguar from BZ, 6 samples per jaguar from CEX

BZ: samples taken three days each week, three samples per day. NEX: 6 days per week, three samples per day

BZ: highest cortisol on higher visitor days, NEX: no difference

Speothos venaticus

Belo Horizonte Zoo

Locomotion, eating, drinking, resting, sleeping, vigilance, vocalisation, positive interaction, negative interaction, abnormal, depart from public, approach public, OOS

Observations four times per day. Used instantaneuous recording with scan sampling or focal animal sampling. Each sp observed 172 times.

Number and behaviour

Species from closed habitat presented an increase in the frequency of eating in the presence of large public groups and an increase in positive interactions in the presence of medium public category compared to open habitat species. On the contrary, the alert behaviour presented a decrease expressed by the closed habitat species, in comparison to open habitat ones, in the presence of medium and large public groups. In comparison to diurnal species, the nocturnal ones ate less, were more alert, and moved less away from the visitors when large public groups were around. Moreover, still in comparison to diurnal animals, the nocturnal species expressed fewer positive interactions and move less in the direction of visitors when a medium public category was present. The carnivore species moved more, rested less, expressed less positive social interactions, and approached more the visitors when medium visitor groups were around, and were more alert and moved more away from visitors when large visitor groups were present. The omnivorous mammals ate more and rested more in the presence of large visitor groups. Omnivorous animals decrease the positive interaction when medium visitor group was present and interacted more when large visitor groups were around. Arboreal and scansorial species presented the same response pattern in the presence of large visitor groups eating more, drinking less, and resting less. In addition, the alert behaviour was less expressed by arboreal and scansorial species in the presence of medium visitor category and by the scansorial species in the presence of large groups. Arboreal animals interacted more when medium size visitor groups were present and less and large groups were around. And the scansorial species moved more away from visitors and was more non-visible in the presence of large visitor groups

Chrysocyon brachyurus

Leopardus pardalis

Suricata suricatta

140 feacal samples in 10 social groups at 8 zoos. 2.6hrs of data on visitor number captured on day of faecal samples and the day before. 2 min scans during 20 minute observations.

Looked at median and SD visitor numbers on the day of faecal sample and the day before

The median number of visitors at the enclosure was positively correlated with fGC concentrations on the following day, with variation in the visitor numbers having the opposite effect. Groups with consistently high median number of visitors had higher FGMs

Multi institution (n=4)

UK and South Africa

vigilant, positive HAIs, negative HAIs, feeding, comfort, positive social, negative social, locomotion, interaction with the environment, resting, ARBs, vocalising, preening, OOS. Position within enclosure

Behavioural observations undertaken 1 to 4 times per day. Each observation lasted 5 minutes. One minute inter-scan interval. Number observations varied per site from 6 to 86 closed and 12 to 83 open.

Presence/absence

Indoor and outdoor exhibits

SPI for enclosure usage

Meerkats showed increased positive social interactions, increased alert behaviours, and reduced environmental interactions in the first month post-opening, as compared to closure periods. They also used more of their enclosures during periods of closure and spent longer than would be expected in zones furthest from visitor viewing areas when facilities reopened. African penguins showed no behavioural change between open and closure periods. Enclosure usage during both observation periods was relatively even and no differences were observed in enclosure use between open and closure periods.

Panthera tigris altaica

Dierenpark Amersfoort

12 weeks, 1 day per week. 18 observations per day (9 per tiger). 397 faecal samples.

Aggressive, affiliative, stereotypy and inactivity

3 x 5 minute focal observations, 15secs apart per tiger. Three separate observation periods per day. Average of numbers of visitors at the start and end of each 5 minute period

563m2 enclosure

Influence of average visitor number on behaviour

For the female, increased visitor count correlated with decreased activity and increased resting behaviours during morning obs, no impact during the midday slot, in the afternoon decreased rest and increase in activity in relation to visitors. increase in aggressive behaviour was only observed in the Morning time slot. For the male, in the Morning time slot an increase in active behaviours of the male tiger correlated with an increase in visitors. frequency of stereotypic behaviour increased signi cantly with visitor count in the Morning and midday time slot.

Tremarctos ornatus

Cleveland Metroparks Zoo

3yrs 5 months, 45.7 hrs per bear

Visible, stereotypy, social, foraging, locomotion, inactive

Maximum of twice per day, minimum two days per week. 10 minute focal follows, behaviour recorded every 30 seconds

Presence/absence, number, number in zoo

Indoor and outdoor access, with access available to an off exhibit area 364.7m2

Visibility calculated as number of intervals per observation the bear was visible in the habitat, divivded by total number of intervals (n=20). Behaviours calculated as a percentage of visible intervals in which behaviour performed, divided by total visible intervals (i.e. proportion of visible time)

Bears were more visible when guests were present. Bears spent more time in locomotion when larger crowds were present. Bears spent less time inactive when large crowds were present at the habitat compared to when small crowds or no crowds were present, and when no crowds were present compared to small crowds. Inactivity was lower on days of high daily attendance compared to medium attendance. No impact of daily attendance on locomotion or foraging. No impact on stereotypies.

Indoor and outdoor access, with access available to an off exhibit area 364.7m2

Helarctos malayanus

Indoor and outdoor access, with access available to an off exhibit area 364.7m2

Ursus arctos horribilis

Indoor and outdoor access, with access available to an off exhibit area 350m2

Ursus americanus

Indoor and outdoor access, with access available to an off exhibit area 350m2

Canis lupus baileyi

1 x safari park, 2 x zoo

3 weeks (72hrs per wolf)

Behaviour & Physiology

lying, standing, eating, walking, running

Behaviour obs 4 days per week, 6hrs per day. Total of 72hrs per wolf. 5 minute scan sampling. 120 faecal samples collected from 10 wolves (12/wolf)

4 x naturalistic enclosures (sizes: 680, 776, 604 and 790m2)

Less time lying and less time eating when there were more visitors in the zoo. Higher cortisol on Sunday's than Saturday, Monday and Tuesday. Locomotion higher when visitor numbers were lower.

1yr 9 months (123.75 hr per bear)

stereotypical (locomotor), normal active, inactive, and out of sight

15 minute observations with one minute scan sampling. 35.0 hr of observations per bear were made in the fall, 31.0 hr in the winter, 29.8 hr in the spring, and 28.0 hr in the summer

4423m2 outdoor enclosure

Behaviour calculated as a proportion of 'in sight' time

One bear increased stereotypies and reduced time spent inactive with bigger crowds, two bears reduced stereotypies. One increased time spent inactive. The other did not change period of time spent inactive.

4 days per week per observation phase (24 observation days total). Faecal samples two - three times per week for banteng and five or more times per week for grizzly and polar bears.

Behaviour & Physiology

Behaviour - space use; Physiological - FGMs

4 x 6 week phases of observations. 2 videos 2hrs in length recorded per day. 3 minute scan sampling to calculate number of animals in each enclosure zone.

143 - 156 samples per animal (31 - 41 samples per animal per phase). 73 - 92.6 hours per phase (1466 1851 observation points)

The grizzly bears spent more time in their habitat than in the den when visitors returned. No change in FGMs

Behaviour & Physiology

143 - 156 samples per animal (31 - 41 samples per animal per phase). 83 - 87 hours per phase (1676 to 1739 observation points per phase)

Polar bear spent more time near viewing areas after visitors returned. No change in FGMs.

Leptailurus serval

Werribee Open Range Zoo

Manipulated exposure to visitors through changing encounters

servals alternated between four different treatments, involving interactive presentations - typically high visitor numbers, behind-the-scenes encounters, both activities combined, or no interaction at al

Behaviour & Physiology

Behaviour (passive, active, maintenance, stereotypies), behavioural diversity and FGMs

Weekly blocks (7 days - treatment exposed over the 7 days) of four different treatments were imposed three times on each animal over 12 weeks. Order of treatments was randomised. Observations were 30 - 45 minutes long for each treatment type. Behaviour sampled every 60 seconds within the observation period.

Interaction programme

Off display to the public (apart from during interactions). Solitary housed, swapped between an open yard (75m2) and pens (36m2)

four daily recording sessions per animal over three consecutive days per treatment (the last 3 days of treatment), using instantaneous scan sampling every 60 s. Individual faecal samples were collected daily. Observations undertaken before, during and after the encounter (plus also in the morning and afternoon).

Reduction in stereotypic pacing on weeks when participating in interactive presentations, or the two activities combined. Behavioural diversity reduced on weeks when cats participated in both activities. Adrenocortical activity did not vary between treatments.

Zalophus californianus

Six Flags Discovery Kingdom

Behaviour (play alone, social play, maintenance, tactile, feeding, water rest, land rest, pattern swim, random swim, move on land, scan on land, scan in water, aggression, nursing/suckling, object obstruction), enclosure use

Two focal follows using continuous sampling, each 7.5 min long, were conducted per day, for each individual. Maximum number of visitors was recorded during each focal follow.

Presence/absence

SPI to investigate enclosure use

No impact of visitors on behaviour greater preference for the water bordering visitor viewing areas when visitors present

More time submerged and more time alert in the water when visitors were present, and less time engaged in social interactions. More time feeding when visitors present. Increased use of water areas near visitor viewing areas when visitors present.

Alert (rest with head up, stand, walk, run, stalk, vocalisation), rest

4 x 2hr sessions, 2 minute scan sampling

Number and noise

Increased alert with increased numbers, increased alert with increased noise

vigilance, aggression, stereotypic, rest

5 x 2hr sessions, 1 minute scans

percent scans at each abundance level

Increased vigilance

vigilance (scan, approach, track, follow), rest

5 x 2hr sessions, 2 minute scans

Increased vigilance, reduced rest

Panthera pardus orientalis

Belo Horizonte Zoo

20hrs (10hrs with visitors, 10 hrs without)

Behaviour, enclosure use, sound (dB)

Scan sampling with a 2 minute interval for 20 minute periods

Presence/absence and noise

Calculated behaviour as a percentage per observation session

No difference in behaviour or enclosure use

Leopardus pardalis

Speothos venaticus

4 months (two months per transition period - one month with/without visitors)

Behaviour & Physiology

FGMs, behaviour: charge glass, glass strike, hiss, OOS, environmental interaction, stereotypy, social interaction, locomotion, groom, stationary

before and after covid closure and before and after reopening. Faecal samples collected three times per week during the two transition periods. 15 samples per open/closure period per transition period (1 and 2). Behaviour: one min scan sampling over 20 min observation periods twice per day 2 days per week. Open data from observation period 1 was based on 60 min observation periods.

Presence/absence

3 x indoor off exhibit areas (109ft, 80ft, 80ft) and an on show grass paddock (19,200ft)

Data calculated as % of time to make 60 min and 20min obs comparable

(1) significant increases in fGM concentrations for cheetahs and giraffes between the transition periods but not within them; (2) a significant increase in time spent ‘not visible’ in the cheetahs in the second transition period; and (3) increased vigilance behaviors in the giraffes after the zoos closure

Ailuropoda melanoleuca

5 months (72hrs per panda)

Decibels recorded every 10 mins and averaged per hour, visitors were total number of people per hour

Number and noise

Indoor and outdoor enclosures

No impact of sound levels, duration of door directed behaviour higher when more people

Panthera tigris altaica

Inactive, explore, locomotion, affiliative, self-groom, pacing, off show plus location in enclosure

Continuous focal sampling, 30 minute observations. Visitor number, decibels and tiger location undertaken every 3 minutes and then averaged for the 30 minute time period. 80 x 30 minute observations.

Number and noise

Outside (2749m2) and inside (126m2)

SPI for enclosure use

No impact of visitor noise or presence on tiger behaviour

Pathera tigris altaica

active, resting, sleep, abnormal, OOS

3 x 2 day period of observations - 2 days 2 weeks nefpre the eent, 2 days during the event, 2 days one week after the event. 90 - 93 observations per species. 60 minute intervals on data collection - each hr observed walked pre defined route. Observations undertaken from 12:00 - to 23:00. Presence/absence of behaviour within the grp recorded.

Looked at relationship between decibels at the enclosure (dBA and dBC) and behaviour, and also event period (pre, during, post)

More active and less likely to rest with higher dBC but less active with higher dBA

More active with higher dBC

More active and less resting with higher dBA. More likely to be resting during than pre-event.

Arctictis binturong

More resting at higher dBA

More likely to be OOS with higher dBA

Panthera pardus orientalis

More likely to be OOS with higher noise (dBC)

Less likely to be asleep at higher dbA

Leopardus pardalis

More abnormal behaviour at lower dBA. Less likelto rest post event than during the events, and more likely to show abnormal behaviour pre event than during

Lontra Canadensis

65 days over 21 months

10 minute observation blocks with 30 second intervals (scan and one-zero sampling). Av 85 mins of observation per day. Total observations 10,108 minutes

Number and behaviour

naturalistic exhibit

Visitor number unrelated to ARBs, more ARBs seen when visitor effect score higher

Suricata suricatta

72hrs (36hrs per treatment) over 8 days (4 days unregulated, 4 days regulated)

Regulation of visitor behaviour

Use of signage requesting visitors do not ineract with the animals and be generally quiet

Behaviour (sentry, vigilance, locomotion, foraging, social interaction, sleeping, OOS) and enclosure use (orientation to visitor viewing area)

2minute observations in 6 x 30 minute observation blocks.

Number, noise, behaviour

Open air, enriched outdoor exhibits. All animals had access to off show dens.

average proportion of time spent engaged in each observed behaviour, the average time spent looking towards visitors and the average proximity to the viewing area (m) on each study day

No effect of treatment on prxomity to viewing areas or meerkat behaviour

9 (5 visitor presence/absence at fence; 4 encounter programme)

on display, 5M. During encounter 4 U

Cheetah Outreach

On display 28 - 30.5hrs. During encounters 1.97 - 4.52hrs, 21 - 63 encounters. HR data available for 0.12 to 0.98hrs during 4 - 11 encountrers

Behaviour & Physiology

Behaviour: whilst on display: active - pacing, observing, climbing, jumping, running, walking, auto grooming, allogrooming, crouching, tail flciking, purring; inactive - lying head up, lying head down plus location in enclosure; behaviour during encounters - purring, tail flicking, lying with head down and HR

On display: 1 minute scan sampling for 2 - 3 hrs per day. During encounter: 15 second scan sampling

Interaction programme, number

On display - behaviour categorised as abnormal or inactive. During encounters: proportion of observations in different behaviours during and between encounters

On display: pacing lower when visitors present, no difference in inactive or tail flicking behaviour. No difference in enclosure use. In encounters: no difference in tail flicking or purring when stroked. Increased time lying with head down when stroked. No impact of number of people. HR lower when stroked. No relationship between HR and number of people.

Arctictis binturong whitei

184.6hrs (n=5 days) open, 163.2hrs (n=6 days) closed

Active (den use, travel, vigilance), inactive

Indoor (103m2) and outdoor (2290m2) areas

Analysed data during 24hr periods and during zoo opening times

No difference in den use, travel, vigilance either during the full 24hr comparison nor during opening hours only

Ursus arctos arctos

8, 11 and 14 closed days at the three zoos (and the same open)

4 behaviour categories: activity (locomotion, foraging, engagement with enrichment, self directed behaviour), inactivity, abnormal activity (begging, pacing), and out of sight, and location in enclosure

6 to 10 scans per day, recording behaviour and location in enclosure

65 to 300m2, 2 x zoos gave access to outside and inside, 1 x zoo only outside). Zoos with ad lib inside access only had public viewing on one side. Zoos with no inside access had public viewing on two sides.

SPI for enclosure use. Behaviour compared between open and closed periods.

Zoo 1: decreased the time spent active due to the increased time they spent on begging; Zoo 2: no difference in activity budget between conditions; Zoo 3: less time inactive after facility reopening. At two of the zoos (those where public can only access from one side and they can go inside), zone use was more equal when open to the public - using both inside and outside enclosures. Bear at MZ spent less time inside and more time close to the public when it reopened. During the Zoo Closed conditions, the YZ female spent 66.7% of her time in Zone 1 (closest to visitors). During the Zoo Open conditions, the SPI index significantly decreased. The bear began spending significantly more time in Zone 3 (farthest from visitors) and decreased her time spent in Zone 1. No difference in the other zoo (where visitors have access at two sides and they cannot go inside).

6 - 10 hrs per day over 6 days

Active, resting, stereotypic

5 min scan sampling for a period of 6-10 hrs. 2 days in on-exhibit area (visitors present), 2 days off-exhibit area, 2 days no visitors

Presence/absence

16 x on exhibit (outside), 2 x off exhibit (inside)

proportion of time spent exhibiting each behavior. Compared visitor presence (2 days on exhibit) vs 2 days vistor absence

Rested more when visitors were present, and were more active on visitor absence days.

National Zoo Park

Active (climbing, cooling, drinking, eating, excreting, grooming, licking, olfaction, playing, rubbing, rolling over, running, scratching, scent marking, vocalisation, and walking), inactive (lying on back, resting awake, sitting, sleeping), stereotypic

Animals observed for 6hrs per day for 4 consecutive days. Focal animal sampling at one minute intervals. Visitor number captured every one minute.

858m2 and 1445m2

Stereotypy increased with increasing visitors

Stereotypy increased with increasing visitors

n=10 zoo, n=2 research centre

Animals not on display at 4 facilities (17M 16F). Faecal samples collected every other day for 6 weeks.

Animals on public display had higher FGMs than those housed off display

Parque Ecológico de São Carlos

80hrs (40hrs with enrichment, 40hrs without)

Stereotypic pacing

Focal animal sampling for 2 - 3 hrs. Duration (minutes) the animal engaged in pacing in each sample period. Total number of visitors in the enclosure per observation period.

128m2 per enclosure

Pacing increased with visitors