First Advisor

Deborah Duffield

Date of Publication

Winter 3-16-2015

Document Type


Degree Name

Doctor of Philosophy (Ph.D.) in Biology






African elephant -- Behavior, Asiatic elephant -- Behavior, Environmental enrichment (Animal culture), Global Positioning System, Accelerometers, Elephants -- Habitat -- Design, Zoo exhibits -- Design



Physical Description

1 online resource (xii, 140 pages)


African savanna elephants (Loxodonta africana) and Asian elephants (Elephas maximus) are a focus of welfare research in zoos due to their high intelligence, complex social structure, and sheer size. Due to these challenges, some argue that zoos are inherently incapable of providing appropriate care for elephants, while others believe that zoos can fulfill the needs of these species with improved husbandry. There is a general consensus from both within and outside of zoos, however, that zoos must improve their elephant programs or cease exhibiting these animals altogether. Now more than ever, applied research on zoo elephant welfare is needed to provide context for this debate.

Researchers are interested in how far zoo elephants walk due to the potential health and welfare benefits of walking in these highly mobile species. Zoo researchers recently adopted GPS technology to study elephant walking, and preliminary evidence suggests that African elephants in large zoo exhibits walk distances that correspond with wild elephants under non-extreme conditions. However, data are limited from Asian elephants and from elephants in more typically-sized exhibits. In Chapter Two, I discuss important methodological considerations of utilizing GPS in a zoo environment, including an introduction to the technology, sources of error and mitigation, methods to improve GPS performance, and possible effects of GPS device attachment on animal behavior. This review shows GPS performance is adequate for tracking zoo elephant walking when proper methodological techniques are applied, and should serve as a useful reference for zoo researchers considering using GPS.

In Chapter Three, I used GPS anklets to measure outdoor daily walking distance in 56 adult female African (n = 33) and Asian (n = 23) elephants housed in 30 zoos. I collected 259 days of data and found that elephants walked an average of 5.34 km/day with no significant difference between species. Multivariate regression models predicted that elephants with more dynamic feeding regimens (more diverse feeding types and frequencies; unscheduled feeding times) will walk more. Distance walked was also predicted to be higher in elephants that spend time in a greater number of different social groups. Distance walked was predicted to decline with age. Finally, I found a significant negative correlation between distance walked and nighttime space experience. The results of the analysis suggest that zoos that want to increase walking in their elephants need not rely solely on larger exhibits, but can increase walking by adding quality and complexity to exhibits. However, my results failed to establish a definitive link between walking distance and other validated measures of elephant welfare. Thus, the direct health and welfare benefits of walking in zoo elephants remain unresolved.

Resting behaviors are an essential component of animal welfare, but have received little attention in zoological research. In Chapter Four, I used accelerometers in anklets to complete the first large-scale multi-species investigation of zoo elephant recumbence. I collected 344 days of data from 72 adult female African (n = 44) and Asian (n = 28) elephants at 40 zoos. I found that African elephants are recumbent an average of 2.14 hours/day, which is significantly less than Asian elephants at 3.22 hours/day. Multivariate regression models predicted that African elephant recumbence increases when they experience more space at night, and Asian elephant recumbence increases when they spend time housed alone. Both species showed a similar response to substrate, such that African elephants spending time on all-hard substrates are predicted to be recumbent less, while Asian elephants spending time on all-soft substrates are predicted to be recumbent more. The discovery that occasional non-recumbence is a common behavior in zoo elephants also introduces a new area of research that may have important animal welfare consequences. Finally, this study established that zoos should continue their efforts to replace hard substrate with soft substrate in order to provide zoo elephants with environments that facilitate recumbence.

Overall, this work assessed walking and recumbence in zoo elephants, which will allow zoos to gauge the prevalence of these behaviors in their elephants as compared to the sub-population studied here. A variety of factors that are associated with these behaviors were also identified. With this information, zoos can prioritize modifications to their facilities and animal management programs to create an environment that encourages zoo elephants to express walking and recumbence behavior, should they choose to do so.

This work is one component of the Elephant Welfare Project, the largest zoo animal welfare project ever undertaken, and is unprecedented in both scope and scale. The project was funded by the Institute of Museum and Library Services (IMLS), an independent, U.S., federal, grant-making agency that supports libraries, museums, and zoos. At the time of this writing, the first manuscripts from this project are being submitted to academic journals. These papers will describe the prevalence and distribution of a variety of elephant behaviors and welfare indicators, serve as a benchmark for future elephant welfare studies, and aid in decision making with regard to best practices in elephant management.


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