Concerns about animal welfare increasingly shape people's views about the acceptability of keeping animals for food production, biomedical research, and in zoos. The field of animal welfare science ...has developed over the past 50 years as a method of investigating these concerns via research that assesses how living in human-controlled environments influences the behavior, health and affective states of animals. Initially, animal welfare research focused on animals in agricultural settings, but the field has expanded to zoos because good animal welfare is essential to zoos' mission of promoting connections between animals and visitors and raising awareness of conservation issues. A particular challenge for zoos is ensuring good animal welfare for long-lived, highly social species like elephants. Our main goal in conducting an epidemiological study of African (Loxodonta africana) and Asian (Elephas maximus) elephant welfare in 68 accredited North American zoos was to understand the prevalence of welfare indicators in the population and determine the aspects of an elephant's zoo environment, social life and management that are most important to prevent and reduce a variety of welfare problems. In this overview, we provide a summary of the findings of the nine papers in the collection titled: Epidemiological Investigations of North American Zoo Elephant Welfare with a focus on the life history, social, housing, and management factors found to be associated with particular aspects of elephant welfare, including the performance of abnormal behavior, foot and joint problems, recumbence, walking rates, and reproductive health issues. Social and management factors were found to be important for multiple indicators of welfare, while exhibit space was found to be less influential than expected. This body of work results from the largest prospective zoo-based animal welfare study conducted to date and sets in motion the process of using science-based welfare benchmarks to optimize care of zoo elephants.
Stereotypic behavior is an important indicator of compromised welfare. Zoo elephants are documented to perform stereotypic behavior, but the factors that contribute to performance have not been ...systematically assessed. We collected behavioral data on 89 elephants (47 African Loxodonta africana, 42 Asian Elephas maximus) at 39 North American zoos during the summer and winter. Elephants were videoed for a median of 12 daytime hours per season. A subset of 32 elephants (19 African, 13 Asian) was also observed live for a median of 10.5 nighttime hours. Percentages of visible behavior scans were calculated from five minute instantaneous samples. Stereotypic behavior was the second most commonly performed behavior (after feeding), making up 15.5% of observations during the daytime and 24.8% at nighttime. Negative binomial regression models fitted with generalized estimating equations were used to determine which social, housing, management, life history, and demographic variables were associated with daytime and nighttime stereotypic behavior rates. Species was a significant risk factor in both models, with Asian elephants at greater risk (daytime: p<0.001, Risk Ratio = 4.087; nighttime: p<0.001, Risk Ratio = 8.015). For both species, spending time housed separately (p<0.001, Risk Ratio = 1.009), and having experienced inter-zoo transfers (p<0.001, Risk Ratio = 1.175), increased the risk of performing higher rates of stereotypy during the day, while spending more time with juvenile elephants (p<0.001, Risk Ratio = 0.985), and engaging with zoo staff reduced this risk (p = 0.018, Risk Ratio = 0.988). At night, spending more time in environments with both indoor and outdoor areas (p = 0.013, Risk Ratio = 0.987) and in larger social groups (p = 0.039, Risk Ratio = 0.752) corresponded with reduced risk of performing higher rates of stereotypy, while having experienced inter-zoo transfers (p = 0.033, Risk Ratio = 1.115) increased this risk. Overall, our results indicate that factors related to the social environment are most influential in predicting elephant stereotypic behavior rates.
Parrots are popular companion animals, but are frequently relinquished because of behavioral problems, including abnormal repetitive behaviors like feather damaging behavior and stereotypy. In ...addition to contributing to pet relinquishment, these behaviors are important as potential indicators of diminished psychological well-being. While abnormal behaviors are common in captive animals, their presence and/or severity varies between animals of the same species that are experiencing the same environmental conditions. Personality differences could contribute to this observed individual variation, as they are known risk factors for stress sensitivity and affective disorders in humans. The goal of this study was to assess the relationship between personality and the development and severity of abnormal behaviors in captive-bred orange-winged Amazon parrots (Amazona amazonica). We monitored between-individual behavioral differences in enrichment-reared parrots of known personality types before, during, and after enrichment deprivation. We predicted that parrots with higher scores for neurotic-like personality traits would be more susceptible to enrichment deprivation and develop more abnormal behaviors. Our results partially supported this hypothesis, but also showed that distinct personality dimensions were related to different forms of abnormal behavior. While neuroticism-like traits were linked to feather damaging behavior, extraversion-like traits were negatively related to stereotypic behavior. More extraverted birds showed resiliency to environmental stress, developing fewer stereotypies during enrichment deprivation and showing lower levels of these behaviors following re-enrichment. Our data, together with the results of the few studies conducted on other species, suggest that, as in humans, certain personality types render individual animals more susceptible or resilient to environmental stress. Further, this susceptibility/resiliency can have a long-term effect on behavior, as evidenced by behavioral changes that persisted despite re-enrichment. Ours is the first study evaluating the relationship between personality dimensions, environment, and abnormal behaviors in an avian species.
We evaluated 255 African (Loxodonta africana) and Asian (Elephas maximus) elephants living in 68 North American zoos over one year to quantify housing and social variables. All parameters were ...quantified for the both the day and the night and comparisons were made across these time periods as well as by species and sex. To assess housing, we evaluated not only total exhibit size, but also individual animals' experiences based on the time they spent in the unique environments into which the exhibits were subdivided. Variables developed to assess housing included measurements of area as a function of time (Total Space Experience), environment type (Indoor, Outdoor, In/Out Choice) and time spent on hard and soft flooring. Over the year, Total Space Experience values ranged from 1,273 square feet to 169,692 square feet, with Day values significantly greater than Night values (p<0.001). Elephants spent an average of 55.1% of their time outdoors, 28.9% indoors, and 16% in areas with a choice between being in or out. Time spent on hard flooring substrate ranged from 0% to 66.7%, with Night values significantly greater than Day (p<0.001). Social factors included number of animals functionally housed together (Social Experience) and social group characteristics such as time spent with juveniles and in mixed-sex groups. Overall Social Experience scores ranged from 1 to 11.2 and were significantly greater during the Day than at Night (p<0.001). There were few significant social or housing differences between African (N = 138) and Asian (N = 117) species or between males (N = 54) and females (N = 201). The most notable exception was Total Space Experience, with African and male elephants having larger Total Space Experience than Asian and female elephants, respectively (P-value<0.05). The housing and social variables evaluated herein have been used in a series of subsequent epidemiological analyses relating to various elephant welfare outcomes.
This study quantified social life events hypothesized to affect the welfare of zoo African and Asian elephants, focusing on animals that were part of a large multi-disciplinary, multi-institutional ...elephant welfare study in North America. Age was calculated based on recorded birth dates and an age-based account of life event data for each elephant was compiled. These event histories included facility transfers, births and deaths of offspring, and births and deaths of non-offspring herd mates. Each event was evaluated as a total number of events per elephant, lifetime rate of event exposure, and age at first event exposure. These were then compared across three categories: species (African vs. Asian); sex (male vs. female); and origin (imported vs. captive-born). Mean age distributions differed (p<0.05) between the categories: African elephants were 6 years younger than Asian elephants, males were 12 years younger than females, and captive-born elephants were 20 years younger than imported elephants. Overall, the number of transfers ranged from 0 to 10, with a 33% higher age-adjusted transfer rate for imported African than imported Asian elephants, and 37% lower rate for imported females than males (p<0.05). Other differences (p<0.05) included a 96% higher rate of offspring births for captive-born females than those imported from range countries, a 159% higher rate of birthing event exposures for captive-born males than for their imported counterparts, and Asian elephant females being 4 years younger than African females when they produced their first calf. In summarizing demographic and social life events of elephants in North American zoos, we found both qualitative and quantitative differences in the early lives of imported versus captive-born elephants that could have long-term welfare implications.
•Elephants spent less time and were more variable in expressing tactile contact than proximity.•Keepers had excellent levels of agreement in rating elephant bond strength.•More time and consistency ...in proximity by elephants predicted “strong or strongest bond” ratings by keepers.
Understanding the affiliative social relationships, or bonds, between zoo elephants has implications for both their welfare and management, yet there is limited work assessing and describing these bonds. Consequently, there is a need for the development of a reliable assessment tool. We used multiple metrics of proximity and tactile contact, as well as keeper surveys, to assess the social bond strength of 41 elephant dyads from 22 different zoos. Survey descriptions of social bond strength were based on previous research and included proximity and separation-reunion behaviors between individuals in a dyad. Approximately half of the elephant dyads in our study were rated as having a “strong” or “strongest” bond by keepers, who showed excellent agreement in their ratings of elephant bond strength (ICC(1,k)=0.82). Elephant dyads that spent more time in proximity (within two elephant body lengths), and those that were more consistent in this behavior across time had an increased predicted probability of being rated as having a “strong or strongest bond” by keepers (p<0.001; p=0.002; respectively). Affiliative tactile contact within dyads, described using duration, diversity, symmetry, and variability metrics, was not significantly related to keeper assessments of dyad bond strength. On average, proximity within dyads occurred more often and was less variable than dyads’ tactile behaviors. Our results suggest that tactile contact may play a more limited role in the maintenance of zoo elephant social bonds than proximity; however, additional research is needed to confirm this. Additionally, this study suggests that keepers are accurately assessing the proximity behavior of their elephants, highlighting the potential of this survey tool to reliably measure the social bond strength of zoo elephant dyads.
The prevalence of stable behavioural differences between individuals of a species (i.e. personality) is puzzling because it indicates that there are limits on animals' behavioural plasticity and, ...therefore, optimality of behaviour. Personality may result from individual state characteristics (e.g. morphology or physiology). In turn, these characteristics can lead to differential fitness outcomes for individuals. Cognitive processing of environmental information may be such a characteristic. We developed a subjective personality assessment for Amazona amazonica. We then assessed whether personality predicted a cognitive state difference in attention bias, as measured by the proportion of balks and errors when performing a spatial foraging task in the presence of a passive human observer. Attention biases occur either because individuals attend more quickly to certain environmental stimuli, or because they cannot disengage their attention from such stimuli. Two factors, ‘neuroticism’ and ‘extraversion’, accounted for 66% of the total variance in personality. There was individual variation between parrots' scores on both personality factors and both factors were temporally consistent over 1 year. There was a significant correlation between neuroticism and attention bias. Evolutionarily, attention biases are selected for because the fitness cost of failing to attend to potential threats is much greater than the cost of expending energy attending to benign stimuli. Therefore, cognitive biases such as attention bias are logical candidate cognitive states driving stable personality differences. Our findings show that differences in personality in A. amazonica are correlated with attention bias, a biologically relevant difference in cognition.
•Temporally stable personality factors exist in Amazona amazonica.•These factors can be reliably measured by multiple raters, and predict behaviour.•Our results provide experimental support for a link between cognition and personality.•We found that more neurotic parrots showed greater attention bias for environmental stimuli.•Attention biases were negatively correlated with performance on a foraging task.
The management of African (Loxodonta africana) and Asian (Elephas maximus) elephants in zoos involves a range of practices including feeding, exercise, training, and environmental enrichment. These ...practices are necessary to meet the elephants' nutritional, healthcare, and husbandry needs. However, these practices are not standardized, resulting in likely variation among zoos as well as differences in the way they are applied to individual elephants within a zoo. To characterize elephant management in North America, we collected survey data from zoos accredited by the Association of Zoos and Aquariums, developed 26 variables, generated population level descriptive statistics, and analyzed them to identify differences attributable to sex and species. Sixty-seven zoos submitted surveys describing the management of 224 elephants and the training experiences of 227 elephants. Asian elephants spent more time managed (defined as interacting directly with staff) than Africans (mean time managed: Asians = 56.9%; Africans = 48.6%; p<0.001), and managed time increased by 20.2% for every year of age for both species. Enrichment, feeding, and exercise programs were evaluated using diversity indices, with mean scores across zoos in the midrange for these measures. There were an average of 7.2 feedings every 24-hour period, with only 1.2 occurring during the nighttime. Feeding schedules were predictable at 47.5% of zoos. We also calculated the relative use of rewarding and aversive techniques employed during training interactions. The population median was seven on a scale from one (representing only aversive stimuli) to nine (representing only rewarding stimuli). The results of our study provide essential information for understanding management variation that could be relevant to welfare. Furthermore, the variables we created have been used in subsequent elephant welfare analyses.
Cognitive mechanisms are an important part of the organization of the behavior systems of animals. In the wild, animals regularly face problems that they must overcome in order to survive and thrive. ...Solving such problems often requires animals to process, store, retrieve, and act upon information from the environment—in other words, to use their cognitive skills. For example, animals may have to use navigational, tool-making or cooperative social skills in order to procure their food. However, many enrichment programs for captive animals do not include the integration of these types of cognitive challenges. Thus, foraging enrichments typically are designed to facilitate the physical expression of feeding behaviors such as food-searching and food consumption, but not to facilitate complex problem solving behaviors related to food acquisition. Challenging animals by presenting them with problems is almost certainly a source of frustration and stress. However, we suggest here that this is an important, and even necessary, feature of an enrichment program, as long as animals also possess the skills and resources to effectively solve the problems with which they are presented. We discuss this with reference to theories about the emotional consequences of coping with challenge, the association between lack of challenge and the development of abnormal behavior, and the benefits of stress (arousal) in facilitating learning and memory of relevant skills. Much remains to be done to provide empirical support for these theories. However, they do point the way to a practical approach to improving animal welfare—to design enrichments to facilitate the cognitive mechanisms which underlie the performance of complex behaviors that cannot be performed due to the restrictions inherent to the captive environment.
Two states in the United States now have legislation requiring that laying hens be provided with sufficient space to perform particular behaviors. To provide a framework for translating these ...performance standards into a space requirement, kinematic analysis was used to measure the amount of space needed for White Leghorn hens to stand, turn around 180°, lie down, and wing flap. Hyline W-36 hens (n = 9) were marked on the tops of their heads and the tips of both wings and 3 toes with black livestock marker. Each hen was then placed in a floor pen (91.4 × 91.4 cm) and filmed using 2 high-speed cameras. The resulting images were processed using a software program that generated 3-dimensional space use for each behavior. Because none of the hens lay down in the test pen, the 2-dimensional space required for lying was determined by superimposing a grid over videos of the hens lying down in their home cages. On average, hens required a mean area of 563 (± 8) cm(2) to stand, 1,316 (± 23) cm(2) to turn around, 318 (± 6) cm(2) to lie down, and 1,693 (± 136) cm(2) to wing flap. The mean heights used were 34.8 (± 1.3) cm for standing, 38.6 (± 2.3) cm for turning, and 49.5 (± 1.8) cm for wing flapping. However, space requirements for hens housed in multiple-hen groups in cage or noncage systems cannot be based simply on information about the space required for local movement by a single hen. It must also incorporate consideration of the tendency of hens in a flock to synchronize their behaviors. In addition, it must include not just local movement space but also the space that hens may need to use for longer-distance movements to access resources such as food, water, perches, and nest boxes.
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