The effects of visitor density and intensity on the behavior of two captive jaguars (Panthera onca)

Several researchers have reported significant effects of visitor density and intensity on captive animal behavior. This study determined whether this was the case for 2 captive jaguars housed at the Woodland Park Zoo, Seattle, WA. Subjects were monitored for changes in behavior as a function of visitor density and intensity. The jaguars were observed for 8 hr per week for 29 weeks—March 31 until October 11, 1998—for a total of 230 hr. Continuous frequency sampling was used, and visitor density and intensity were recorded every minute. Parametric statistics were used to test for correlations between behavior and density, intensity, or a combination of the two. Both density and intensity were significant for time spent non-visible for both cats, and intensity showed a significant effect on the female's pacing behavior. In addition, the male cat exhibited a trend for increased aggression based on both visitor density and intensity and a trend of intensity affecting his social behavior. In conclusion, both density and intensity had a significant effect on behavior, with intensity showing a larger effect.     

Visitors' Effects on the Welfare of Animals in the Zoo: A Review

Since the 1970s, research about zoo visitors' effects on the welfare of nonhuman animals in captivity has intensified. Numerous studies have shown that characteristics such as visitor presence, density, activity, size, and position are associated with animal behavioral and—to alesser extent physiological—changes. Studies usually interpret these changes as negative (undesirable) or positive (enriching), but it remains unclear whether they significantly impinge on animal welfare. To make confident conclusions about visitors' effects necessitates more studies using (a) a wider range of animal groupings, (b) measures of stress, (c) visitor-animal variables, and (d) other methodological improvements In the meantime, in addition to further research, individual zoos need to emphasize (a) monitoring the stress indicators of their captive animals, (b) observing visitor behavior, and (c) ensuring that staffs are aware of the “visitor effect” concept.     

The “Visitor Effect” and captive black-tailed prairie dog behavior

The presence and density of people at a zoo exhibit is associated with behavioral changes in captive animals. However, most of these studies have only investigated behavioral changes in primates. To widen the range of animal groups studied, we examined the effects of visitor observation on captive black-tailed prairie dog (Cynomys ludovicianus) behavior and enclosure distribution. Due to the rich variety of readily-observable, sophisticated behaviors the black-tailed prairie dog is a wonderful model species for the study of behavior. In this study, we were able to quantify the effects of observation on key behaviors.     

Individual Effects of Seasonal Changes,Visitor Density,and Concurrent Bear Behavior on Stereotypical Behaviors in Captive Polar Bears (Ursus maritimus)

Stereotypical behaviors in captive polar bears (Ursus maritimus) can be detrimental to their welfare. These behaviors can be reduced through enrichment programs but are often not completely eliminated, so identifying potential triggers is important. The present study investigated the influences of seasonal changes, visitor density, and concurrent bear activity on stereotypical behaviors exhibited by 3 captive polar bears at the Toronto Zoo. All bears exhibited these behaviors; however, individual differences were found in duration and form. The male exhibited less stereotypical behavior during spring, and the females exhibited less stereotypical behavior during winter. An increase in visitor density was associated with more stereotypical behavior in 1 female but less stereotypical behavior in the other 2 bears. All bears engaged in more stereotypical behaviors when the other bears were inactive, and 1 female engaged in more stereotypical behaviors when the other bears were out of sight. Further, when conspecifics were active, all bears engaged in less stereotypical behaviors. Given the variability among individual bears, future enrichment programs must be tailored to the needs of individuals to maximize efficacy.     

Visitor effects on the behavior of captive Western lowland gorillas: the importance of individual differences in examining welfare

Given the conflicting and somewhat limited findings available on the effect of zoo visitors on primate behavior, the primary purpose of this study was to provide additional data on gorillas' response to variations in crowd size and to look at what other factors—both intrinsic (e.g. personality, sex, and rearing history) and extrinsic (e.g. group)—might influence such responses. Subjects included four groups (three mixed‐sex and one bachelor) of captive western lowland gorillas living at Zoo Atlanta. Overall activity budgets and rates of social behaviors in high and low crowd conditions were compared. Behaviors suggestive of general welfare did not vary as a function of crowd size alone, although we did observe one significant interaction and several trends. Specifically, two groups spent more time engaged in undesirable behaviors in the high crowd condition, whereas the other two groups showed the opposite response pattern. Additionally, males, but not females, showed a trend toward greater rates of contact aggression in the high crowd condition, although wounding rates did not vary with crowd size. We also found trends towards variation in response to crowd size as a function of individual personality ratings. These results suggest that although the visitor effect may constitute an impediment to optimal animal welfare, this may only apply to some individuals or groups. We emphasize the importance of continuing to explore individual differences and the limitations of a one‐size‐fits‐all approach when describing influences on animal welfare. Zoo Biol 31:586‐599, 2012. © 2011 Wiley Periodicals, Inc.     

Retrospective study of morbidity and mortality of captive jaguars (Panthera onca) in North America: 1982–2002

One hundred seventy‐two medical records of captive jaguars (Panthera onca) were examined from 30 American Zoo and Aquarium Association (AZA) institutions housing jaguars between 1982–2002. The study determined common causes of morbidity and mortality, and the influence of age, gender, and melengestrol‐acetate (MGA) exposure on these rates. The most common causes of morbidity in captive jaguars were found to be dental, gastrointestinal, integumentary, and musculoskeletal diseases. Prevalence of types of disease varied with age, with older animals experiencing a higher prevalence of multiple types of disease. Females developed reproductive disease more frequently than males, and the data suggest that MGA exposure increased the risk of developing female reproductive disease. The most common causes of mortality were reproductive diseases in females and musculoskeletal diseases in males. There was a high rate of neonate and pediatric mortality, primarily due to stillbirths or unexplained neonatal death, trauma, and pneumonia. Other diseases or clinical signs that seemed remarkable were a high prevalence of episodes of epistaxis among young, as well as old, jaguars. Based on these findings, management suggestions for the captive jaguar population are presented. Zoo Biol 0:1–12, 2006. © 2006 Wiley‐Liss, Inc.     

游客密度对动物园中黇鹿行为的影响

2006年9月20日-10月10日,以江苏省扬州市茱萸湾公园半散放鲇鹿为研究对象,采用目标动物取样法和全事件记录法,研究了不同游客密度条件下黇鹿日行为时间分配.并探讨了游客密度对鲇鹿日活动的影响.结果表明:在游客高峰期与低峰期,雄性取食、观望行为,雌性观望以及幼鹿取食、观望和修饰行为之间均存在显著差异(P＜0.05).在游客低峰期,雄性与雌性取食、反刍、卧息行为,雄性与幼鹿观望、其它行为,雌性与幼鹿取食、卧息、观望均存在极显著差异(P＜0.01);在游客高峰期,雄性与雌性取食、卧息,雌性与幼鹿取食、卧息、观望及移动行为均差异极显著(P＜0.01),雄性和幼鹿观望行为差异显著(P＜0.05).不同游客密度条件下游客活动对鲇鹿的取食、观望具有显著影响,当游客密度达到6人·min-1以上时,雄性观望行为显著增加,当达到16人·min-1以上时,雄性取食行为差异极显著(P＜0.01),此时雌性和幼鹿观望行为差异亦达到极显著水平(P＜0·01).因此,在旅游管理中,应将游客密度限制在16人·min-1以下,以减少对鲇鹿日活动影响.     

An Hourly Variation in Zoo Visitor Interest: Measurement and Significance for Animal Welfare Research

A methodological difficulty facing welfare research on nonhuman animals in the zoo is the large number of uncontrolled variables due to variation within and between study sites. Zoo visitors act as uncontrolled variables, with number, density, size, and behavior constantly changing. This is worrisome because previous research linked visitor variables to animal behavioral changes indicative of stress. There are implications for research design: Studies not accounting for visitors' effect on animal welfare risk confounding (visitor) variables distorting their findings. Zoos need methods to measure and minimize effects of visitor behavior and to ensure that there are no hidden variables in research models. This article identifies a previously unreported variable—hourly variation (decrease) in visitor interest—that may impinge on animal welfare and validates a methodology for measuring it. That visitor interest wanes across the course of the day has important implications for animal welfare management; visitor effects on animal welfare are likely to occur, or intensify, during the morning or in earlier visits when visitor interest is greatest. This article discusses this issue and possible solutions to reduce visitor effects on animal well-being.     

The effect of visitors on the behavior of zoo-housed western lowland gorillas (Gorilla gorilla gorilla)

Primates, especially apes, are popular with the public, often attracting large crowds. These crowds could cause behavioral change in captive primates, whether positive, neutral, or negative. We examined the impact of visitors on the behavior of six western lowland gorillas (Gorilla gorilla gorilla), observing the troop over 6 weeks during high season (4.5 hr/day, 35 days, May–July 2016). We used focal scan sampling to determine activity budget and enclosure usage, and focal continuous sampling to identify bouts of anxiety-related behavior (visitor-directed vigilance, self-scratching, and aggression). Both daily zoo-entry numbers (V_GATE) and instantaneous crowds at the exhibit (V_DENSITY) were measured. Overall, V_GATE had little effect across behaviors. However, consistent with the more acute time frame of measurement, V_DENSITY was a better predictor of behavior; at high crowd volumes, we observed significant group-level changes in activity budget (increased inactivity, increased locomotion, and decreased environment-related behaviors), increase in some anxiety-related behaviors, and decreased enclosure usage. Although contributing similar effects, it could not be determined if crowd numbers, composition, or noise most affected the troop, nor any chronic effects of exposure to large crowds. Nevertheless, our findings suggest that measures to minimize the impacts of large crowds at the exhibit would be beneficial. Furthermore, we highlight potential discrepancies between common methods for measuring visitor numbers: V_GATE is less sensitive to detecting visitor effects on behavioral indices than V_DENSITY. Future studies should appropriately match the biological time frame of welfare indicators and visitor measures used to ensure the reliability of findings.     

Influence of zoo visitor presence on the behavior of captive Indian gaur (Bos gaurus gaurus) in a zoological park

Visitors to zoos can be a source of potential disturbance and stress to some captive, nonhuman animals in the wild. To determine the influence of visitor presence on captive bison (Bos gaurus gaurus), the study analyzed the behavior of 4 individuals at the Arignar Anna Zoological Park, India. The study often observed the behavior of the animals on visitor-present days and on days when visitors were absent. In the presence of zoo visitors, the bison showed a higher level of intragroup aggression and moving behavior. In contrast, the bison rested more when no visitors were present. The results revealed that the presence of zoo visitors significantly influenced the behavior of captive bison and thereby may have affected their welfare.     

Influence of Zoo Visitor Presence on the Behavior of Captive Indian Gaur (Bos gaurus gaurus) in a Zoological Park

Visitors to zoos can be a source of potential disturbance and stress to some captive, nonhuman animals in the wild. To determine the influence of visitor presence on captive bison (Bos gaurus gaurus), the study analyzed the behavior of 4 individuals at the Arignar Anna Zoological Park, India. The study often observed the behavior of the animals on visitor-present days and on days when visitors were absent. In the presence of zoo visitors, the bison showed a higher level of intragroup aggression and moving behavior. In contrast, the bison rested more when no visitors were present. The results revealed that the presence of zoo visitors significantly influenced the behavior of captive bison and thereby may have affected their welfare.     

Jaguar and Puma Activity Patterns and Predator‐Prey Interactions in Four Brazilian Biomes

Jaguars (Panthera onca) and pumas (Puma concolor) coexist throughout the Neotropics. Using camera trapping in four Brazilian biomes, we compare the daily activity patterns of the jaguar and puma, and their relationships with their main prey species. We used a kernel density method to quantify daily activity patterns and to investigate overlap between these predators and their main prey. Both cats showed intensive nocturnal and crepuscular activity (0.69 and 0.14 kernel density, respectively, for jaguars; 0.68 and 0.19 kernel density, respectively, for pumas). Only in the Pantanal did we observe a pattern of concentrated diurnal activity for both species. We found little temporal segregation between jaguars and pumas, as they showed similar activity patterns with high coefficients of overlapping (average ?₁ = 0.86; SE = 0.15). We also observed a significant overlap between the activity patterns of the predators and their main prey species, suggesting that both predators adjust their activity to reduce their foraging energy expenditure. Our findings suggest that temporal partitioning is probably not a generalized mechanism of coexistence between jaguars and pumas; instead, the partitioning of habitat/space use and food resources may play a larger role in mediating top predator coexistence. Knowledge about these behavior aspects is crucial to elucidating the factors that enable coexistence of jaguars and pumas. Furthermore, an understanding of their respective activity periods is relevant to management and associated research efforts.     

Activity budget and spatial distribution of Bennett's wallabies (Macropus rufogriseus) in open versus closed exhibit designs

Although many studies investigating the impacts of zoo exhibit designs on captive animals exist, none have been performed on how they influence the behavior and welfare of captive Bennett's wallabies (Macropus rufogriseus). Here, we assess the impact of exhibit design on the activity budget and spatial distribution of Bennett's wallabies. We compared animal behavior in two open exhibits (i.e. physical interaction between animals and visitors permitted) to two closed exhibits (i.e. physical interaction between animals and visitors prohibited). Behavioral data were collected using focal sampling, and spatial distribution was recorded on exhibit maps at regular time intervals. We found a significant increase in feeding and interactive behaviors in closed exhibits in comparison to open exhibits. However, other behaviors such as resting, locomotion, and vigilance did not vary with design. Functional use of space was similar between both designs; however, the effect of habituation may be relevant to consider in future studies. Although some support for visitor effects were present, our study provided no evidence for strong impacts of exhibit design on Bennett's wallaby welfare. Our study emphasizes the need for additional research into the impacts of how zoo environments affect Bennett's wallaby behavior and welfare.     

The Effect of Fission–Fusion Zoo Housing on Hormonal and Behavioral Indicators of Stress in Bornean Orangutans (Pongo pygmaeus)

The welfare of captive animals could be improved if zoos were to place more emphasis on their species-specific needs. In the wild, orangutans live in a fission–fusion social system and have a semisolitary lifestyle. However, most zoos keep orangutans in permanent groups, which may be stressful for them. Apenheul Primate Park in Apeldoorn, Netherlands, houses 14 Bornean orangutans in a simulated fission–fusion social system. To assess how this housing system affects indicators of stress, we measured fecal glucocorticoid metabolites (fGCM) and determined proportions of self-directed behavior (scratching and autogrooming). We compared fGCM concentrations of the Apenheul animals to those of zoo orangutans kept in permanent groups. In addition, we tested the effect of group size, visitor number, sex, age, and change of group composition on fGCM concentrations and proportions of self-directed behavior in the Apenheul orangutans. In contrast to Bornean orangutans housed in permanent groups, we did not find a group size effect on fGCM or on self-directed behavior in Apenheul’s fission–fusion housing system. In addition, fGCM concentrations in Apenheul orangutans increased significantly with visitor numbers. Visitor number also affected proportions of self-directed behavior, and mean proportions of scratching were positively correlated with mean values of fGCM concentrations. Although these results suggest that the fission–fusion housing system in Apenheul reduces the group size effect leading to social stress in Bornean orangutans, they also show that visitors are an important factor that needs to be mitigated if the well-being of captive primates and other zoo animals is to be improved.     

Visitors' effects on the welfare of animals in the zoo: a review

Since the 1970s, research about zoo visitors' effects on the welfare of nonhuman animals in captivity has intensified. Numerous studies have shown that characteristics such as visitor presence, density, activity, size, and position are associated with animal behavioral and—to alesser extent physiological—changes. Studies usually interpret these changes as negative (undesirable) or positive (enriching), but it remains unclear whether they significantly impinge on animal welfare. To make confident conclusions about visitors' effects necessitates more studies using (a) a wider range of animal groupings, (b) measures of stress, (c) visitor-animal variables, and (d) other methodological improvements In the meantime, in addition to further research, individual zoos need to emphasize (a) monitoring the stress indicators of their captive animals, (b) observing visitor behavior, and (c) ensuring that staffs are aware of the “visitor effect” concept.     

Establishment of a Captive All‐male Group of Proboscis Monkey (Nasalis larvatus) at the Singapore Zoo

Surplus male proboscis monkeys at the Singapore Zoo pose a considerable problem for maintenance and maximizing of exhibition potential. In 2008, a new exhibit was constructed to house and display a group of six proboscis monkey males born in Singapore Zoo. To document and monitor the all‐male group establishment in the new exhibit, we conducted observations on intragroup interactions between the monkeys, spatial use of their new exhibit, and visitor effects on their behavior. We found contact aggressive interactions between the monkeys to be consistently lower than noncontact aggressive interactions and by week six of introduction to the new exhibit, contact aggression was almost nonevident. Affiliative interactions also developed between individuals in the group, with an interface of aggressive and socioreconcilatory behavior influenced by food competition and a dominance hierarchy. This was evident from significantly higher overall aggression and affiliation during feeding times compared to nonfeeding times, and this was reduced when food competition was mitigated by modifying the feeding regime. We measured the groups’ spatial use of the exhibit and the relation to behavior, crowd size, and density. Our results showed that the proboscis monkeys utilized the available exhibit space, were largely unaffected by visitor crowd size and density, and were able to exhibit a variety of natural behaviors, including swimming. Our accomplishment in maintaining and displaying an all‐male group of proboscis monkeys in captivity provides viable options for more comprehensive captive management and breeding programs for this endangered species. Zoo Biol. 32:281‐290, 2013. © 2012 Wiley Periodicals, Inc.     

Use of enclosure space by captive lion-tailed macaques (Macaca silenus) housed in Indian zoos

Captive nonhuman animals use enclosure space differentially. Enclosure features strongly influence this. This study recorded both the enclosure space used by 47 captive lion-tailed macaques housed in 13 zoos across India and the behavior of the macaques. The exhibition of abnormal behaviors, food-related behaviors, and social interactions correlated significantly with the use of the edge zone (the part of the enclosure closest to the visitor area). Animals housed in barren enclosures used the edge zone to a significantly greater percentage than did those housed in complex exhibits. Percentages of autogrooming, social interactions, and food-related behaviors significantly correlated with the use of the enrich zone. Space use studies assist in recognizing areas within the enclosure, which captive animals actively use. Conversely, the studies can identify areas infrequently used and show how to make maximum use of these enclosure areas. Further studies targeting both the increase in percentages of natural behaviors exhibited and use of the enrich zone used the current study on captive lion-tailed macaques for their design.     

Infanticide in a jaguar (<Emphasis Type="Italic">Panthera onca</Emphasis>) population—does the provision of livestock carcasses increase the risk?

Infanticide is an antagonistic behavior that may provide an evolutionary benefit for the perpetrator. Cases of infanticide have rarely been reported in Neotropical carnivores. The objective of this study was to provide empirical evidence of infanticide in a local jaguar (Panthera onca) population in the Brazilian Pantanal. We present infanticide data from opportunistic sampling. Each month, from 2013 to 2015, we monitored pastures for evidence of livestock carcasses. A remotely triggered camera trap was placed at encountered fresh cattle carcasses. Through this monitoring, two cases of infanticide were recorded. Both cases occurred in close proximity to a cattle carcass, a large prey item that may attract multiple jaguars—including females with cubs around age of weaning—and increase the risk of infanticide. Our observations provide new empirical evidence of jaguar social behavior and intraspecific competitive interactions, potentially driven by locally high population density and resource provision.     

Frustrated Appetitive Foraging Behavior,Stereotypic Pacing,and Fecal Glucocorticoid Levels in Snow Leopards (Uncia uncia) in the Zurich Zoo

One common visitor complaint in zoos is that the nonhuman animals are not visible. This problem needs to be resolved without compromising the animals' welfare; environmental enrichment could solve the problem. This study investigated whether enrichment would increase public exposure time of lowland tapir (Tapirus terrestris) in the Belo Horizonte Zoo in Minas Gerais, Brazil. Observations were made before (62 hr) and during (62 hr) the introduction of enrichment using focal animal sampling with instantaneous recording of behavior. The 5 enrichment items were a bamboo fence covered in vines, logs, a sandbox, dry leaves, and bamboo bushes. Before the enrichments were applied, the tapir was not visible to the public for more than 85% of the time. In addition, during the analysis of the enrichment treatment, other variables were considered—such as weekday, time of day, and weather conditions—which could influence the animals' interaction with the enrichments. The enrichments increased and decreased the expression of some behaviors; however, public viewing time of the animals did not increase. Thus, the enrichment applied was not strong enough to overcome the animals' crepuscular behavior.     

Conflict and reconciliation behavior trends of the bottlenose dolphin (Tursiops truncatus)

Wild bottlenose dolphin (Tursiops truncatus) populations display societal structures characterized by numerous and frequent changes in group composition, complex social relationships, and high levels of cooperation, attributes also observed in human and nonhuman primate cultures. Maintaining social relationships under such elemental conditions can frequently create conflict—and the opportunity for reconciliation—among group members. The conflict and reconciliation behavior patterns of the chimpanzee (Pan troglodytes) have been studied extensively; trends are well‐documented in the wild and in captivity. Apparent cultural similarities have prompted several analogous studies on wild and captive bottlenose dolphins. This research attempted to corroborate previous efforts by analyzing the social behavior of seven captive bottlenose dolphins to determine the effects of sex and age on the frequency of conflict and reconciliation, and to investigate the incidence of consolatory behavior within the group. A total of 3,428 interactions involving focal animals, 414 of which were conflict episodes, were documented during 261 hr of videotaped observations. Although the sample size precluded meaningful statistical evaluation of the influence of sex on conflict and reconciliation, participant age was a significant determinant of conflict frequency. Conversely, age did not impact frequency of reconciliation, which only occurred after 18% of all conflict interactions. Little to no definitive evidence of consolation was apparent within the study group. While results partially support the findings of previous dolphin reconciliation research, extensive behavioral studies of wild populations should be conducted before generating broad comparisons between human terms and nonhuman behavioral interactions. Zoo Biol 29:567–585, 2010. © 2009 Wiley‐Liss, Inc.