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Humans are a constant in the lives of captive animals, but the effects of human-animal interactions vary. Research on the welfare impacts of human-animal interactions focus predominantly on mammals, whereas fish have been overlooked. To address this lack of research, we assessed the impacts of aquarium visitors on the behaviors of ten members of four elasmobranch species: an Atlantic stingray (Dasyatis sabina), four southern stingrays (Hypanus americanus), two blue-spotted maskrays (Neotrygon kuhlii), and three fiddler rays (Trygonorrhina dumerilii). The rays engaged in a significantly higher proportion of active behaviors and a lower proportion of inactive behaviors when visitor density levels were high; however, there were no significant changes for negative or social behaviors. Individual analyses indicated that all three fiddler rays and one of the southern stingrays' active behaviors differed across visitor density levels, whereas there was no association between active behavior and visitor density levels for the other rays. Further research is needed to determine whether this pattern is an adaptive or maladaptive response to visitors, but this research provides much needed initial data on activity budgets within elasmobranch species.

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Visitors are a prominent feature in the lives of zoo animals, and their presence can cause a range of impacts on zoo animals (typically classed as positive, negative or neutral impacts), commonly referred to as the 'visitor effect'. This paper quantitatively collates the literature on the visitor effect in non-primate species, investigates the types of measures used to assess impacts of visitors on animals and considers whether impacts vary across non-primate species in zoos. In total, there were 105 papers which had investigated the impact of zoo visitors on 252 non-primate species/species groups. There has been a steady increase in visitor effect research in zoos since 2012 and this body of work incorporates species from avian (28% study species), reptilian (9%), amphibian (2%), fish (4%) and invertebrate taxa (1%). However, there is still a bias towards mammalian species (56%). The response to visitors varied across taxa. Amphibians responded negatively to visitors more frequently than would be expected by chance (p < 0.05), birds responded neutrally more frequently than would be expected by chance (p < 0.05) and fish responded neutrally and 'unknown' more frequently than would be expected by chance (p < 0.05). This review highlighted a number of animal-based metrics which have been used to assess the impacts of visitors on animals, with measures used varying across taxa. Moving forwards, it is recommended that moving forwards researchers incorporate a suite of measures, incorporating those which are meaningful in terms of being representative of individual animal experiences and animal welfare, collected in a manner which should capture those metrics accurately.

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The potential impact of human presence on captive animal behavior has recently been the focus of considerable research interest, especially following 2020 and 2021 periods of enforced closure as a result of COVID-19 opening restrictions. It is important to investigate whether human presence represents an enriching or stressful stimulus to a range of zoo-housed species. In 2020, during an easing of lockdown restrictions, investigations of the "visitor effect" and "observer effect" were carried out, using the giant otter (Pteronura brasiliensis) as a model species. To investigate the impact of both visitor and observer presence, otter behavior and space use was recorded for a pair of on-show and a pair of off-show otters. Observations were conducted using either a human observer, or cameras, allowing the researchers to investigate otter behavior when no one was present at the exhibits. The Electivity Index was used to assess the otters' use of four enclosure zones. Overall, otter behavior was significantly impacted by observer presence, though the impact of an observer differed between individual otters. Visitors had a minimal effect on otter enclosure use, whereas observers had a greater impact, whereby otters used their pools less frequently and houses more frequently when observers were present. However, this change in zone use differed between individuals, with more dominant otters tending to make use of indoor zones more often when observers were present. Zoos should consider the potential impact of human presence on their animals and use both behavior and space use when conducting their investigations.

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Interest in the impact of human presence on the behavior and well-being of zoo and aquarium animals is increasing. Previous work has conceptualized the presence of zoo visitors as having one of three impacts on the behavior of animals in zoos: positive, negative, or neutral. Research suggests the same species may exhibit all three responses under different conditions, calling into question whether the positive/negative/neutral framework is the most useful way of considering visitor impact on animal behavior. Here we present a model of visitor effects that unifies these three predictions. Our model suggests that zoo-goers may provide a "dither effect" for some animals living in zoos. We posit animals may show nonlinear behavioral responses over a range of visitor densities, effectively exhibiting changes in both comfortable and anxiety-like behaviors under different levels of human presence. We tested this model during two COVID-19 related closures at the San Francisco Zoo, studying seven species for evidence of nonlinear relationships between visitor numbers and animal behavior. Our results support the dither effect acting in several species observed.

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In the zoo, the sound environment experienced by captive wild animals will contain numerous anthropogenic features that may elicit different responses to those stimulated by naturally created, or more biologically relevant, sounds. Husbandry activities, visitor presence and neighbouring species (free-living and captive) will all influence the sounds around zoo-housed species; an animal's behavioural responses may therefore provide an insight into how its welfare state is influenced by this changing sound environment. This project aimed to investigate how animal behaviour was influenced by the sound environment at two large UK zoos; one situated in a more rural location and the other in an urban location. Species were selected based on their location in the zoo, the relevance of sound to their natural ecology (e.g., as a form of communication and/or for anti-predator responses) and their novelty as research subjects in the scientific literature. Behavioural data collection was conducted for five days per enclosure per zoo at the population and individual level for birds housed in different styles of enclosure. Instantaneous sampling at one-minute intervals was used to collect information on state behaviours, assessed using a pre-determined species-specific ethogram. Event behaviours were collected continuously for each observation period. The sound environment around or in the enclosure was recorded continuously during each behavioural recording session using a recorder mounted on a tripod. Results showed a variety of responses to the presence of visitors and potential associated changes to sound around the enclosure with some behaviours being more influenced by the presence of visitors (and increases in the volume of sound) compared to others, e.g., vigilance and vocalisations. Overall, birds showed few of the changes thought to indicate poor or impoverished welfare states linked to changes to the sound environment in their enclosure or to the presence of visitors, but we recommend that zoos consider further measurement and recording of sound on a species-by-species basis to capture individual responses and behavioural changes to variation in visitor number and the sound environment.

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Primates are some of the most cognitively advanced species held in zoos, and their interactions with visitors are complex. The COVID-19 pandemic provided a unique opportunity to understand the impact of zoo visitors on animals, in comparison to "empty zoos". This study sought to understand the impact of facility closures and subsequent reopenings on behavioural and physiological parameters of welfare in four primate species housed in the UK: bonobos (Pan paniscus) (n = 8), chimpanzees (Pan troglodytes) (n = 11), and western lowland gorillas (Gorilla gorilla gorilla) (n = 6) held at Twycross Zoo (TZ); and olive baboons (Papio anubis) (n = 192) held at Knowsley Safari (KS). Behavioural data were collected from April-September 2020 (KS) and November 2020-January 2021 (TZ). Faecal samples were collected during morning checks from October-November (TZ) and July-November 2020 (KS). Faecal glucocorticoid metabolites (FGMs) were measured using ELISA kits. Statistical analysis for behavioural observations was undertaken using general linear models. Enclosure usage was assessed using t-tests and Mann-Whitney U-tests as appropriate. Bonobos and gorillas spent less time alone when facilities were open to the public (p = 0.004, p = 0.02 respectively). Gorillas spent less time resting when the facility was open to the public (p = 0.04), and chimpanzees engaged in more feeding (p = 0.02) and engagement with enrichment (p = 0.03) when the zoo was open to the public than when it was closed. Olive baboons performed less sexual and dominance behaviour and approached visitor cars more frequently when the safari park was opened to the public than they did the ranger's vehicle during closure periods. There were no significant changes in physiological parameters for any of the study species. The results suggest variable impacts of the zoo closures on zoo-housed primates. We recommend future work that seeks to understand the impact of individual-level differences on "visitor effects" and that differences between animal experiences in zoos and safari parks are further explored in a range of species.

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The effect of visitor presence on zoo animals has been explored in numerous studies over the past two decades. However, the opportunities for observations without visitors have been very limited at most institutions. In 2020, the Oregon Zoo was closed, in response to the global SARS-CoV-2 (COVID-19) pandemic, from 15 March 2020 to 12 July 2020, resulting in approximately four consecutive months without visitor presence. This study aimed to quantify potential behavioral and hormonal changes expressed during two transition periods in zoo visitor attendance: the initial time period before and after closure in March 2020 and time before and after reopening in July 2020. Fecal glucocorticoid metabolite (fGM) concentrations of resident giraffes (n = 2) and cheetahs (n = 2) were tracked using enzyme immunoassay (EIA) analyses. Average fGM concentrations during the two transition periods were compared using a two-way mixed ANOVA. Additionally, twice-weekly scan sampling was used to quantify behavioral observations across the transitions, which were analyzed as individual behavior proportions. Individual behavior proportions were compared across the Zoo's opening status and time of day using Kruskal-Wallis (H) tests. The results of our analyses showed the following outcomes: (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 immediately after the Zoo's closure. However, the changes observed in fGM concentrations may be more strongly correlated with concomitant social changes (giraffes) and some medical events (cheetahs) rather than with the Zoo's opening status. Nevertheless, this study was able to quantify differences in behavioral frequencies and fGM concentration in cheetahs and giraffes at the Oregon Zoo during the times of transition between visitor's presence and absence. The results indicate that, while there was a possible, but relatively minor impact of the presence and absence of visitors on some behaviors, the differences observed in fGM concentration may have been more affected by some of the concomitant social changes and medical events that happened during the same period than by the presence or absence of visitors.

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This study investigated the behaviour of two Elasmobranch species; Southern fiddler ray (Trygonorrhina dumerilii, n = 1) and Port Jackson shark (Heterodontus portusjacksoni, n = 4) and two teleost species; moonlighter (Tilodon sexfasciatus, n = 1) and banded morwong (Cheilodactylus spectabilis, n = 1) living within a single enclosure. For this study, two treatments were compared, the original enclosure design, and then after the enclosure had been renovated to more closely represent the species natural habitats, with a raised front viewing glass to prevent visitor interaction. Behaviours such as resting, swimming and abnormal behaviours such as surface and perimeter swimming (elasmobranchs only) were recorded as well as location within the enclosure, for 10 days pre and 10 days post renovation. The Port Jackson sharks significantly reduced the performance of abnormal behaviours after renovation, and significantly increased the time spent near the exhibit front. The Southern fiddler ray increased resting post renovation, while the teleost species also spent more time near the exhibit front. Although a small sample size was used, the results suggest that a more naturalistic environment with multiple micro-habitats and effective visitor barriers allows for a greater proportion of the day spent exhibiting natural behaviours, greater space use and reduced stereotypes.

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Amphibians are an understudied group in the zoo-focussed literature. Whilst commonly housed in specialist exhibits and of real conservation value due to the global extinction crisis, amphibian welfare is not often investigated empirically in zoo settings. The limited research that is available suggests that enclosure design (structure, planting and naturalistic theming) has a positive impact on the time that amphibians will be on show to visitors. However, the categorisation of any "visitor effect" (i.e., influences of visitor presence on amphibian activity and time on display) is hard to find. The COVID-19 pandemic forced the closure of zoological organisations in the UK for several months from March 2020, with gradual re-openings from the summer into autumn and winter. This event provided a unique opportunity to study the effect of the lack of visitors, the presence of essential zoo staff only, the wider return of organisational staff, and then the return of visitors over a prolonged period. This project at WWT Slimbridge Wetlands Centre assessed the number of individuals of six species of amphibian-common toad (Bufo bufo), common frog (Rana temporaria), smooth newt (Lissotriton vulgaris), pool frog (Pelophylax lessonae), golden mantella (Mantella aurantiaca) and golden poison dart frog (Phyllobates terribilis)-visible to observers under different conditions. All amphibians were housed in a purpose-built indoor exhibit of individual enclosures and were recorded when visible (as a proportion of the total population of the enclosure) during closure, the return of extra centre staff and visitor periods. The results showed species-specific differences in visibility, with some species of amphibian being more likely to be on view when the presence of people at their enclosure was less likely or in smaller numbers. Such differences are likely related to the specific camouflage or anti-predation tactics in these focal species. Further study to quantify amphibian sensitivity to, and perception of, environmental change caused by public presence (e.g., light levels and sound) would be useful welfare-themed research extensions. Our results can help inform husbandry, collection planning and amphibian enclosure design to reduce any noticeable visitor effects, and provide a useful benchmark for further, more complex, welfare assessment measures.

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The sound environment of a zoo animal is a complex milieu of animal and human-generated sounds; coming from the species itself, other species, visitors, keepers and other zoo-users. Research determining how different components of the sound environment affect animal behaviour is surprisingly lacking but could have real-world impacts for animal welfare and zoo enclosure design. The current study investigated the effects of the sound environment on two flocks of flamingos housed in open-air enclosures at British zoos. Measures of how each flock used its enclosure (as a response variable) and environmental variables (Inband Power and Peak Frequency were recorded as characteristics of the sound environment, as well as temperature, humidity and cloud cover, and finally visitor presence-all as potential predictor variables) were made over a 2-month period. Assessment of space use by zoo animals is often used as a measure of the appropriateness of an exhibit and to understand welfare. Given that flamingo activity is influenced by weather and that the sound environment of the zoo is likely to be influenced by the number and the presence of visitors, it was assumed that these predictor variables would influence where the flamingos were located at different times of the day. As expected, there was a complicated relationship between enclosure use and Inband Power (average spectral density, a measure of sound energy) in both flocks; visitors generated salient sound but other visitor characteristics such as their physical presence may have impacted the movement of the birds around their enclosures. Results show a complex picture where environmental conditions influence flamingo enclosure usage as well as visitor presence and sounds around/in the enclosure. Findings are not consistent between the two flocks, with one flock demonstrating distinct temporal change to enclosure zone occupancy and the other responsive to humidity and cloud cover variation. We believe enclosure use can provide a valuable indication of how birds react to their soundscape; however, our findings suggest more work is needed to unpick the components of captive sound environments, and their relative effects on how animals use their space.

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There is evidence that zoo visitor presence can influence the behaviour and, in some cases, adrenal response of zoo animals, and can sometimes compromise animal welfare. In some laboratory studies, significantly more primate births have been reported on weekends, when fewer people are working there, compared with weekdays when staffing levels are at their highest. Here, we investigate whether there is evidence of a "weekend effect" on births in zoo animals as a result of visitor numbers. Unlike laboratories, zoos are typically busier with visitors on weekends than on weekdays, although staffing levels remain fairly consistent across days of the week. If zoo animal parturition is sensitive to human presence, then fewer births would be expected on weekends compared with weekdays. We tested this using birth data and visitor numbers on the entrance gate from zoo records across 16 species representing artiodactyls, perissodactyls, carnivores and primates at four British zoos, to see whether there is an association between mean daily birth rates and average visitor numbers. We predict that, if there is a visitor effect, daily births should be lower on weekends than weekdays and should correlate with mean daily visitor numbers. Results showed that births for all 16 species were randomly distributed through the week, and there was no significant decline in births on weekends. We conclude that the "weekend effect", if such a thing exists, does not appear to be a feature of zoo births, suggesting that elevated weekend visitor numbers are not sufficiently stressful to trigger delayed parturition.

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The human factor is one of the major determinants of animal welfare in zoos. Assessing changes in activity patterns, behavior and space use due to human presence might help understanding visitors' influence on these animal welfare indicators. In Córdoba (Argentina), we assessed the activity pattern and behavior of Tamandua tetradactyla (lesser anteaters; n = 5) during natural light/dark phases and the animals' use of space. We analyzed responses of lesser anteaters to humans in days open to the public (Sundays and Tuesdays) and days closed to the public (Mondays), collecting data at 5-minute intervals during 12 days, for each animal. Data were analyzed at two temporal scales: a 24 h period and visiting hours. Multivariate analyses showed no differences in activity pattern, behavior and space use among days, exhibiting a consistent response of each individual over the studied days. Principal Component Analysis showed differences between female and male behaviors. Based on these results, the visitor effect on these lesser anteaters would appear to be neutral, indicating that the presence of visitors is at most, a minor concern in the housing and management of these individuals in Córdoba Zoo.

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While previous research has focused on the impact of visitors on zoo-housed animals' behavior, here, we evaluated the impact of visitors on the performance of four zoo-housed Japanese macaques (Macaca fuscata) in a cognitive task. The macaques completed a touchscreen-based match-to-sample task in glass-sided booths at the perimeter of their enclosure, adjacent to a visitor viewing area. The task was novel to all macaques at the start of this study but over the 6-month testing period the macaques showed increased accuracy on the task, suggestive of learning. We recorded the number of visitors within the viewing area roughly every 12 trials each macaque completed. We categorized visitor counts as small (0-20), medium (21-40), and large (41-60) crowds and we considered the macaques' response latencies and accuracy by crowd size and study period (first 3 months versus second 3 months). If visitor presence negatively influenced performance, we predicted that macaques' accuracy would decrease but response times would increase with crowd size. We found effects of crowd size and study period on the macaques' accuracy. In the first period, the macaques performed at chance and accuracy did not differ across crowd categories. In the second period, the macaques' accuracy improved as compared to the first period, but their accuracy was mediated by crowd size: the macaques were significantly more accurate in the presence of small crowds than medium or large crowds. The macaques' response latencies also varied by study period and crowd size, but we found no evidence of a response-slowing effect.

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Captive animal welfare is important for establishments that exhibit species for education, conservation, and research. However, captive animals are often exposed to a number of potential stressors, such as visitors and anthropogenic noise. We aimed to identify the impact of a concert series on the behaviour of Fiordland penguins (Eudyptes pachyrhynchus; n = 2), and solitary- (n = 1) or group- (n = 4)-housed collared peccaries (Pecari tajacu). Animal behaviour, visitor density, and visitor behaviour was monitored pre-concert (afternoons; 16:00-19:00), during the concert (evenings; 19:00-21:00), and post-concert (nights; 21:00-00:00) on concert days (penguin n = 7 days; peccary n = 8 days) and in the same periods on days when there was no concert (penguin n = 8 days; peccary n = 6 days). Fiordland penguins spent more time surface swimming and diving in the pool on concert afternoons and evenings (all p < 0.001), more time in the nest on concert nights (p < 0.001), preened less on concert afternoons and nights (p = 0.019), and engaged with their habitat less on concert evenings and nights (p = 0.002) compared to these periods on days without a concert. The group-housed peccaries slept more in the afternoon and evening (p ≤ 0.01) and were more vigilant at night (p = 0.009) on concert days compared to no-concert days. The solitary-housed peccary slept more on concert nights (p = 0.035), rested more frequently across all time periods on concert days (p < 0.001), and used the front of the enclosure more across all concert time periods (p < 0.001) compared to no-concert days. We provide evidence that behaviour was altered on event days; however, we cannot determine the nature of these changes. Further research is needed to understand the impact of music concerts on zoo animal welfare.

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The impact that humans have on zoo animals can vary based on the species of animal, exhibit design, and individual differences in behavioral responses. We independently analyzed data from 10 never-published studies that examined the impact of zoo visitors on zoo animal behavior. Of the 16 species studied, 90.9% of the mammal species and 60.0% of the fish species demonstrated a change in at least one behavior based on zoo visitor abundance or visitor behavior (e.g., noise, solicitation of interactions from zoo animals). In addition, behavioral changes associated with zoo visitors were present in animals housed in exhibits where there was direct contact with zoo visitors, as well as in exhibits where there was indirect contact and no direct contact. Individuals often varied in their behavioral responses, and some individuals appeared to seek out interactions with visitors. Our findings demonstrate that short-term research projects can provide valuable insight into individual animal-level and species-level responses to visitor abundance and visitor behavior in the zoo setting. We recommend that behavioral assessments focus on the analysis of behaviors of individual animals whenever possible, and we recommend that exhibits provide areas that allow for animals to retreat from the public view.

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Studies on the effects of visitors on zoo animals have shown mixed findings and as a result, the manner in which visitors affect zoo animals remains unclear for many species, including a rarely studied taxa such as penguins. Penguins are a common zoo-housed species and have been shown to display huddling, vigilance and avoidance towards zoo visitors which can be indicative of fear. Here, we examined the effects of covering one visitor viewing area window, out of four, on little penguin (Eudyptula minor) behaviours that may be indicative of fear. Two treatments were randomly imposed on different days: (1) The main visitor viewing area window, where most visitor-penguin interactions occurred, was uncovered ('Main window uncovered') and (2) The main visitor viewing area window was covered ('Main window covered'). Penguin numbers and behaviour were recorded near the main visitor viewing area window and the three other visitor viewing area windows, as well as one area not visible to visitors ('Corner' area). Furthermore, visitor numbers and visitor behaviour were recorded at all four visitor viewing area windows. Covering the main visitor viewing area window reduced the proportion of visitors present at this window by about 85% (p < 0.001) and reduced potentially threatening visitor behaviours at this window such as tactile contact with the window, loud vocalisations and sudden movement (p < 0.05). When the main visitor viewing area window was covered, the proportion of penguins present increased by about 25% (p < 0.05), the proportion of visible penguins preening in the water increased by about 180% (p < 0.05) and the proportion of visible penguins vigilant decreased by about 70% (p < 0.05) in the area near this main window. A preference for the Corner area was also found whereby 59% and 49% of penguins were present in this area when the main window was uncovered and covered, respectively. These results provide limited evidence that the little penguins in this exhibit showed an aversion to the area near the main visitor viewing area window when it was uncovered based on the increased avoidance and vigilance and decreased preening in the water in this area. This suggests visitors may be fear-provoking for these little penguins. However, it is unclear whether visual contact with visitors per se or other aspects of visitor contact, such as visitor-induced sounds and vibrations, were responsible for this apparent aversion when this window was uncovered.

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A "visitor effect" on zoo-housed species has been documented since the 1970s, with research focused on mammals (specifically primates). To broaden our understanding of the "visitor effect" in a non-mammal, we conducted a case study on a pair of hornbills, recording behavior and aviary use alongside of visitor and keeper presence. Temperature and humidity were significant predictors of visitor number, and temperature was a better predictor of hornbill exhibit use than visitor presence. Behavior was significantly affected by the presence of keepers and individual variation in behavior was noted too. Visitor number mediated any interest in a keeper by birds: high visitor number decreased a bird's interest in its keeper. Whilst only a case study on a pair of birds, our research shows that any "visitor effect" is heavily influenced by other environmental variables and that different categories of human (i.e., visitor, keeper) affect how zoo animals utilize their environment.

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The question of 'if and how captive primates are affected by visitors' has gained increasing attention over the last decades. Although the majority reported undesirable effects on behavior and wellbeing, many studies reported contradicting results. Most of these studies were conducted at zoos, typically with little or no control over visitors' actions. Yet little is known about the impact under very controlled visitor conditions. In order to fill this gap, we conducted this study at a primate sanctuary which allows public access only via a guided visit under strict supervision. We observed 14 chimpanzees, recording their behavior during, after and in the absence of guided visits over a 10-month period. Furthermore, we categorized the visitors regarding group size and composition to see if certain group types would produce a stronger impact on the chimpanzees' behavior. As expected, we found visitors at the sanctuary to produce only a neutral impact on the chimpanzees' behavior, detecting a slight increase of locomotion and decrease of inactivity during visitor activities with chimpanzees demonstrating more interest towards larger sized groups. We argue that the impact has been greatly mitigated by the strict visitor restrictions and care strategies allowing chimpanzees a certain control regarding their visibility.

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We examined the effects of regulating the viewing proximity of visitors and the intensity of visitor behaviour on behaviours indicative of fear and stress physiology of 15 zoo-housed little penguins (Eudyptula minor). A 2 × 2 factorial fully randomised design was used to examine the effects of regulating: 1) the viewing proximity of visitors to enclosure, 'normal viewing distance' and 'increased viewing distance' (using a physical barrier set up 2 m away from the enclosure), and 2) the intensity of visitor behaviour, 'unregulated visitor behaviour' and 'regulated visitor behaviour' (using signage and uniformed personnel). In addition, a treatment consisting of closing the enclosure to visitors was included. Penguin behaviour, visitor numbers and visitor behaviour were recorded by CCTV video recordings and direct observations, respectively. Penguin faecal glucocorticoid metabolites were also analysed as a measure of stress physiology. We found that increased viewing distance reduced (p < 0.05) all visitor behaviours except for loud vocalisations and tactile contact with penguins. However, there were no direct effects of signage and uniformed personnel on visitor behaviour (p > 0.05). As the regulation of viewing proximity increased from a closed exhibit to an open exhibit with increased viewing distance, and then to an open exhibit with normal viewing distance, this increased the proportion of penguins huddling (p = 0.0011), vigilant (p = 0.0060) and retreating (p = 0.00013), and decreased the proportion of penguins within 1 m of the visitor viewing area (p = 0.00066), surface swimming (p = 0.00091) and preening in the water (p = 0.042). There were also limited effects of regulating visitor behaviour on penguin behaviour. No treatment effects were found on faecal glucocorticoid metabolites (p > 0.05). These results indicate that regulating visitor viewing proximity affects penguin behaviours indicative of fear and visitor behaviour. This suggests that close visitor contact can be fear-provoking for little penguins but increasing the distance between visitors and penguins can reduce fear responses of penguins by regulating both viewing proximity and visitor behaviour. However, it is unclear whether these changes in penguin behaviour are due to the increased separation between visitors and penguins and/or specific visitor behaviours associated with close viewing proximity to the enclosure, such as leaning over the enclosure or tactile contact with the pool, which are impeded when visitors are further away.

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Diel activity rhythms in mammals are regulated by an endogenous (circadian) timing system which is synchronized by environmental 24-hr periodicities called zeitgebers. Additional direct responses to stochastic environmental factors ensure the fine-tuning to the actual situation and may mask the circadian time course. Following an observational study on behavioral effects of visitor activities in a group of spider monkeys (Ateles geoffroyi) kept free-ranging on a small island of Lake Catemaco, Veracruz, Mexico, we analyzed the effect of weekly varying numbers of visiting tourist boats on the monkeys' diel activity rhythm. With small accelerometer-data logger devices we recorded the monkeys' locomotor activity continuously for several months each. Then we compared the data with those from spider monkeys living without tourist contact. Neither the duration of the monkeys' activity time (α) nor its phase relationship to the 24-hr solar day did change on different weekdays in either site. However, their activity level showed a clear 7-day rhythm. The monkeys of the tourist site showed highest activity on Saturday and Sunday, when the frequency of visiting tourist boats was highest, whereas those of the non-tourist site were least active on Sunday and Monday, when human activities were lowest there. While the monkeys of the non-tourist site usually displayed a distinct bimodal activity pattern peaking in the morning and late afternoon, the pattern in those of the tourist site mostly lacked a morning peak and varied more over time. Based on our results, we suggest that circadian entrainment is not involved in the differences between the diel activity rhythms of the spider monkeys from the two keeping sites and the differing 7-day variation in their activity level. Rather, these differences seemingly reflect direct responses to the differing human activities and thus may correspond to circadian masking effects.

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