Rates of reinforcement and measures of compliance in free and protected contact elephant management systems.

Protected contact is an alternative to traditional captive elephant training techniques that emerged as a result of concerns for animal welfare and personnel safety. The present study documented the behavior of elephants and their animal care professionals to determine rates of reinforcement and measures of compliance under two handling systems. Behavioral data were collected from animal care professionals and elephants during the elephants' baths in both free contact (FC) and protected contact (PC). Positive reinforcement, in the form of food, was delivered, on average, nearly eight times more frequently in the PC condition. Further, the mean rate at which the animal care professionals used the ankus in the FC condition as negative reinforcement was similar to the mean rate at which they provided positive reinforcement to the elephants in the FC condition. Latencies between verbal commands and the elephants' behaviors demonstrated an inconsistent pattern, but were generally longer in the PC condition. The mean percent of "refusals" by the elephants was higher for most behaviors across elephants in the PC condition. The findings suggest that animal care professionals did not heavily rely on positive reinforcement in the FC condition to elicit desired behaviors from the elephants, but this was the case in the PC condition. We propose that longer latencies and higher mean percent of refusals by the elephants may indicate that they were exercising choice or control over their environment, which has been associated with improved well-being. Additional studies of this kind are needed to enable other institutions to make informed decisions about elephant management and welfare.

Advancing Behavior Analysis in Zoos and Aquariums.

Zoos, aquariums, and other captive animal facilities offer promising opportunities to advance the science and practice of behavior analysis. Zoos and aquariums are necessarily concerned with the health and well-being of their charges and are held to a high standard by their supporters (visitors, members, and donors), organized critics, and the media. Zoos and aquariums offer unique venues for teaching and research and a locus for expanding the footprint of behavior analysis. In North America, Europe, and the UK, formal agreements between zoos, aquariums, and university graduate departments have been operating successfully for decades. To expand on this model, it will be necessary to help zoo and aquarium managers throughout the world to recognize the value of behavior analysis in the delivery of essential animal health and welfare services. Academic institutions, administrators, and invested faculty should consider the utility of training students to meet the growing needs of applied behavior analysis in zoos and aquariums and other animal facilities such as primate research centers, sanctuaries, and rescue centers.

Can black-and-white ruffed lemurs (Varecia variegata) solve object permanence tasks?

We examined object permanence in black-and-white-ruffed lemurs (Varecia variegata) at Zoo Atlanta. A series of visible and invisible displacement tasks with suitable controls were presented to five adult subjects. Subjects performed significantly above chance on all regular tasks, except for the double invisible displacements. Subjects failed visible and invisible controls. Failure on the control trials did not appear to be because subjects used the "last box touched" strategy (subjects did not choose the last box touched significantly more than expected by chance). However, a substantial percentage of choices was made to the last box touched by the experimenter. There was no significant difference between this percentage, and the percentage of choices made to the baited box (on both visible and invisible controls), which indicates that subjects were drawn to both boxes which the experimenter visited/touched, and thus failed the controls. Based on the results from the present study, we believe that there is no evidence that black-and-white ruffed lemurs understand visible and invisible tasks in the traditional object permanence battery.

Technology at the zoo: the influence of a touchscreen computer on orangutans and zoo visitors.

A computer-controlled touchscreen apparatus (hereafter referred to as "touchscreen") in the orangutan exhibit at Zoo Atlanta provides enrichment to the animals and allows cognitive research to take place on exhibit. This study investigated the impact of the touchscreen on orangutan behavior and visibility, as well as its impact on zoo visitors. Despite previous research suggesting that providing a single computer system may negatively affect orangutan behavior, there was not a significant increase in aggression, stereotypic, or distress-related behaviors following the activation of the on-exhibit touchscreen. We also investigated the possibility that zoo visitors may be negatively affected by technology because it deviates from naturalism. However, we did not find a change in stay time or overall experience rating when the computer was turned on. This research was the first to assess visitor attitudes toward technology at the zoo, and we found that visitors report highly positive attitudes about technology for both animals and visitors. If subjects visited the exhibit when the computer was turned on, they more strongly agreed that orangutans benefit from interacting with computerized enrichment. This study is the first investigation of an on-exhibit touchscreen in group-housed apes; our findings of no negative effects on the animals or zoo visitors and positive attitudes toward technology suggest a significant value of this practice.

Sex differences in spatial ability: a test of the range size hypothesis in the order Carnivora.

Sex differences in spatial cognition have been reported for many species ranging from voles to humans. The range size hypothesis predicts that sex differences in spatial ability will only occur in species in which the mating system selects for differential range size. Consistent with this prediction, we observed sex differences in spatial ability in giant pandas, a promiscuous species in which males inhabit larger ranges than females, but did not observe sex differences in Asian small-clawed otters, a related monogamous species in which males and females share home ranges. These results provide the first evidence of sex differences in spatial ability in the order Carnivora, and are consistent with the range size hypothesis.

Factors affecting aggression in a captive flock of Chilean flamingos (Phoenicopterus chilensis).

The influence of pair bond status, age and sex on aggression rates in a flock of 84 captive Chilean flamingos at Zoo Atlanta was examined. Analysis showed no difference between aggression rates of male and female flamingos, but adult flamingos had higher rates of aggression than juveniles. There were also significant differences in aggression depending on pair bond status (single, same-sex pair, male-female pair or group). Bonded birds were significantly more aggressive than single birds, which is consistent with the concept that unpaired birds are not breeding and do not need to protect pair bonds or eggs. Birds in typical pair bonds (male-female) and atypical pair bonds (same-sex pairs or groups) exhibited similar rates of aggression. These results contribute to the existing body of research on aggression in captive flamingos.

Factors facilitating research: a survey of zoo and aquarium professionals.

The purpose of this survey was to assess the background, training, and perceptions of professionals conducting and coordinating research at North American zoos and aquariums accredited by the Association of Zoos and Aquariums (AZA) and to identify the factors they considered critical to successful scientific programs. We analyzed responses to a 57-item questionnaire from 231 professionals at AZA zoos and aquariums. The majority of those surveyed conducted behavioral research, conducted research only in a captive setting, held curatorial positions, had their salaries supported by their institutions' operating budget, and considered themselves part of a successful scientific program. About 30% of those we surveyed possessed a doctoral research degree in comparison to 55% possessing lesser level degrees-19% with master's, 34% with bachelor's, and 2% with other degrees. Support from the chief executive officer and personnel dedicated to conducting scientific programs were judged as the two most important factors contributing to the successful scientific programs. The information provided in this report may be used to develop and improve both established and newly initiated scientific programs in zoos and aquariums.

Spatial Design of Guest Feeding Programs and Their Effects on Giraffe Participation and Social Interactions.

Guest-animal feeding programs (GFPs) in zoological institutions aim to foster human-animal connections. The growing establishment of animal welfare science emphasizes the assessment of GFPs as permanent environmental inputs to habitats that require analysis of behavioral output. This study assessed the role of space allocation on giraffe participation and interactions in GFPs in two Florida zoos.Analysis of social structure indicates that centrality and influence from affiliative network on exhibit shape sharing interactions at GFPs under varying management protocols and designs. Findings suggest that interactions and significant ties among conspecifics are context and potentially temporally dependent. We propose the use of multiple guest engagement stations to maximize space and facilitate feeding opportunities for central and peripheral members. This proposed shift would provide variable feeding opportunities that represent group composition following ecological theory while maximizing points of guest engagement. Results indicate that increasing space allocation for GFPs increased feeding bout length for individuals, percent of time sharing among conspecifics, and reduced average rate of displacement/minute. Cross-institutional comparisons indicate that guest programs with more space allocation have lower rates of conspecific displacement.

Spatial memory recall in the giant panda (Ailuropoda melanoleuca).

The current study tested spatial memory recall in 1 male and 1 female giant panda (Ailuropoda melanoleuca). The task required subjects to make a delayed response to a previously lighted location, with varying lengths of delay between the observation phase and the test phase. The male subject reached criterion at 2-, 3-, 4-, 5-, 6-, and 10-s delays. The female subject reached criterion at 2-, 3-, 4-, 5-, 6-, 10-, and 15-s delays. The results support the hypothesis that giant pandas demonstrate significant working memory for spatial location in the absence of external cues, which may be an important mechanism for survival in the wild.

Habituation and desensitization as methods for reducing fearful behavior in singly housed rhesus macaques.

Operant conditioning using positive reinforcement techniques has been used extensively in the management of nonhuman primates in both zoological and laboratory settings. This research project was intended to test the usefulness of counter-conditioning techniques in reducing the fear-responses of singly housed male rhesus macaques living in the laboratory environment. A total of 18 male rhesus macaques (Macaca mulatta) were selected for this project and randomly assigned to one of three groups: a desensitization training group, a husbandry training group, or a control group. Behavioral data were collected before and after a 6 weeks training and/or habituation period during which the first two groups received a total of 125 min of positive reinforcement training (and also were assumed to undergo habituation to the environment) and the control group experienced only simple habituation to the environment. Based on a Wilcoxon Matched-Pairs Sign Test, we found that a significant proportion of animals exposed to desensitization training showed a reduction in the rate at which they engaged in cringing toward humans (exact significance=0.016, one-tailed, N-ties=6), cringing in general (exact significance=0.016, one-tailed, N-ties=6), and in stress-related behaviors (exact significance=0.016, one-tailed, N-ties=6). This was not the case for animals exposed to basic husbandry training or animals in the control group. A significant proportion of desensitization-exposed animals also showed a reduction in the duration of time spent cringing toward humans (exact significance=0.016, one-tailed, N-ties=6), but not in cringing behaviors in general or in stress-related behaviors. There were not a significant proportion of animals in either the husbandry training group or the control group that showed a decrease in duration of these behaviors. Results of this study could enhance both laboratory animal welfare and laboratory animal research, and could be a first step in developing techniques for reducing fearful behavior in rhesus monkeys in the laboratory environment.

Developing a Metric of Usable Space for Zoo Exhibits.

The size of animal exhibits has important effects on their lives and welfare. However, most references to exhibit size only consider floor space and height dimensions, without considering the space afforded by usable features within the exhibit. In this paper, we develop two possible methods for measuring the usable space of zoo exhibits and apply these to a sample exhibit. Having a metric for usable space in place will provide a better reflection of the quality of different exhibits, and enhance comparisons between exhibits.

Twenty-five years of Zoo Biology: a publication analysis.

To expand on earlier publication analyses of the journal Zoo Biology, information from 991 articles published within the journal's first 25 years was compiled and evaluated. Most articles were applied, descriptive accounts that included inferential statistics and/or biological analyses. In addition, most articles focused on captive animals and concentrated on the behavior and reproduction of mammals, with particular emphasis on primates. With respect to authors, the majority of first authors were affiliated with zoos or universities within the United States and collaborated with co-authors at different institutions. Nearly two-fifths of articles acknowledged receiving external funding and external funding increased over the years, however, collaboration did not significantly change receipt of external funding. In addition, first authors with zoo affiliations published more descriptive, applied science articles and acknowledged receipt of external funding significantly less often than first authors with university affiliations. Publication latency was comparable to other scientific journals and increased significantly over the 25-year publication period. Longitudinal trends indicated that experimental, applied science, diet and nutrition, and collaborative articles also increased significantly, whereas descriptive, basic science, behavior, and population biology articles decreased significantly. By describing historical trends in zoo research through an analysis of papers published in Zoo Biology, this article may help to bridge gaps and identify new opportunities. Zoo Biol 27:444-457, 2008. (c) 2008 Wiley-Liss, Inc.

Achieving optimal welfare for the Nile hippopotamus (Hippopotamus amphibius) in North American zoos and aquariums.

Compared to other megafauna managed in zoos and aquariums, the current state of welfare for the Nile hippopotamus (Hippopotamus amphibius) is poorly understood. Complex behavior and physiological characteristics make hippos a difficult species to manage. Thus, hippos in managed care are currently at risk for a decreased state of welfare. In an effort to assess and improve conditions for this species, a survey was administered to North American institutions housing Nile hippos. This assessment utilized a multiple-choice format and consisted of questions relating to group structure, behavior, and exhibit design, allowing for the creation of cross-institutional, welfare-based analysis. Responses were gathered from 85.29% of the institutions to which the survey was distributed. Despite recommendations for maintaining groups of at least five individuals (Forthman, 1998), only 34.25% of hippos in North America were housed in groups of three or more. The survey also highlighted that 39.29% of institutions secure their hippos in holding areas overnight, despite their highly active nocturnal propensities. A better understanding of hippo behavior and environmental preferences can be used to inform wellness-oriented management practices to achieve a state of "optimal welfare".

Relative numerousness judgment and summation in young, middle-aged, and older adult orangutans (Pongo pygmaeus abelii and Pongo pygmaeus pygmaeus).

The ability to select the larger of two quantities ranging from 1 to 5 (relative numerousness judgment [RNJ[) and the ability to select the larger of two pairs of quantities with each pair ranging from 1 to 8 (summation) were evaluated in young, middle-aged, and older adult orangutans (7 Pongo pygmaeus abelii and 2 Pongo pygmaeus pygmaeus). Summation accuracy and RNJ were similar to those of previous reports in apes; however, the pattern of age-related differences with regard to these tasks was different from that previously reported in gorillas. Older orangutans were less accurate than the young and middle-aged for RNJ, and summation accuracy was equivalent among age groups. Evidence was found to suggest that the young and middle-aged based their selection of the largest quantity pair on both quantities within each pair during the summation task. These results show a relationship between subject age and the quantitative abilities of adult orangutans.

The structure of social relationships among captive female giraffe (Giraffa camelopardalis).

Giraffe herds have been characterized as random associations of individuals, but recent evidence suggests giraffe have a more complex social structure. The authors formulated 3 hypotheses designed to evaluate whether a herd of captive giraffe (Giraffa camelopardalis) associated randomly or patterned their behavior and proximity in a manner indicative of social relationships. Affiliative interaction, proximity, and nearest neighbors for 6 captive female giraffe living in a large outdoor enclosure were analyzed, and all three measures were nonrandomly distributed, indicating female giraffe had social preferences. Furthermore, preferences were consistent across measures and time, suggesting that adult female giraffe maintain relationships. Mother-daughter pairs and pairs with large age differences between members interacted and associated most often. The social structure of this captive herd is influenced by social relationships between individual adult females, and the social behavior of individual females should be examined more closely in the wild.

Toward a science of welfare for animals in the zoo.

Although the accredited institutions of the Association of Zoos and Aquariums have all committed to enhancing the welfare of nonhuman animals, acceptable standards and best practices are still under debate. Currently, experts from zoos and the field hold widely divergent opinions about exhibition and management standards for elephants. Standards and practices for managing nonhuman primates provide a model for other nonhuman creatures exhibited in zoos and aquariums. Examining the key issues for primates demonstrates the value of applying scientific data before promulgating standards. The field of applied behavior analysis provides a wealth of information to frame the debate. Animal behaviorists have contributed to an emerging science of animal welfare, which may provide a foundation for empirical zoo management, standards, and practices.

Environmental effects on the behavior of zoo-housed lions and tigers, with a case study of the effects of a visual barrier on pacing.

Tigers and lions in the wild are nocturnal nonhuman animals who may hunt and mate opportunistically during daylight hours. In captivity, they spend most time on exhibit sleeping or pacing. To better understand their activity budget, this study examined the daily behavior patterns of 2 Sumatran tigers and 3 African lions in different housings. The proportion of scans the large felids spent engaged in stereotypic pacing varied by time of day and environment. The tigers spent different amounts of time pacing when housed in different exhibits; the lions paced more in off-exhibit housing than when on exhibit. These differences suggest changes to the cats' immediate housing environment may decrease pacing but provide little insight into altering specifics. Carnivores' pacing relates to their inability to control sensory access to social partners. Both environments with increased pacing contained chain-link fencing. allowing uncontrolled sensory contact. Where the tigers paced, the study placed a visual barrier between one female and keepers' or conspecifics' cues. This did not significantly decrease pacing. However, the study suggests considering sensory access and environmental variables when designing environments for captive carnivores.