The days and nights of zoo elephants: using epidemiology to better understand stereotypic behavior of African elephants (Loxodonta africana) and Asian elephants (Elephas maximus) in north American zoos
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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.
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Elephants typically spend just over 50 percent of their daytime hours under behavioral control in managed activities.
“Elephants typically spend just over 50 percent of their daytime hours under behavioral control in managed activities”
From Continuing challenges of elephant captivity: the captive environment, health issues, and welfare implications (2024) · cites this paper as Greco et al., 2016a
The Discussion (Management Variables) says the studied zoo elephants spent an average of just over 50% of daytime hours in managed activities under a human caretaker’s behavioral control. The claim omits the zoo-elephant setting, so the source does not establish that this is typical of elephants generally. (LLM, conf=0.84)
Experimental — accuracy not yet measuredIn a large-scale epidemiological study of 291 elephants across 70 North American zoos, the Elephant Welfare Initiative underscored the importance of larger social groups for elephant welfare.
“In a large-scale epidemiological study of 291 elephants across 70 North American zoos, the Elephant Welfare Initiative underscored the importance of larger social groups for elephant welfare (Carlstead et al., 2013;Greco et al., 2016a;DiVincenti, McDowell & Herrelko, 2023”
From Continuing challenges of elephant captivity: the captive environment, health issues, and welfare implications (2024) · cites this paper as Greco et al., 2016a
The Discussion (Social Variables; page unavailable) reports that elephants spending more time with larger numbers of herdmates had a lower risk of stereotypic behavior, supporting the welfare point. But the Methods (Data Collection; page unavailable) say this study collected video data from 89 elephants at 39 zoos, not 291 elephants across 70 zoos. (LLM, conf=0.90)
Experimental — accuracy not yet measuredDenying elephants freedoms can result in maladaptation, chronic stress, poor welfare, and abnormal stereotypic behaviors.
“Denying these freedoms can result in maladaptation, chronic stress, poor welfare (32, 33), and abnormal stereotypic behaviors (34, 35).”
From Comparison of animal welfare assessment tools and methodologies: need for an effective approach for captive elephants in Asia (2024) · cites this paper as [35]
There appears to be an inverse relationship between time spent with keepers and an elephant's rate of stereotypy, even outside keeper-directed activities.
“There appears to be an inverse relationship between time spent with keepers and an elephant's rate of stereotypy, even outside of time spent in keeper-directed activities.”
From Continuing challenges of elephant captivity: the captive environment, health issues, and welfare implications (2024) · cites this paper as Greco et al., 2016a
Spending more time with larger numbers of conspecifics and more time with juveniles is associated with reduced stereotypy rates.
“Spending more time with larger numbers of conspecifics and the amount of time spent with juveniles is associated with reduced stereotypy rates”
From Continuing challenges of elephant captivity: the captive environment, health issues, and welfare implications (2024) · cites this paper as Greco et al., 2016a
The social environment has a considerable impact on stereotypic behavior rates, an important indicator of compromised welfare.
“The social environment has a considerable impact on stereotypic behavior rates, an important indicator of compromised welfare”
From Continuing challenges of elephant captivity: the captive environment, health issues, and welfare implications (2024) · cites this paper as Greco et al., 2016a
Being housed separately increases stereotypic behavior.
“being housed separately increases stereotypic behavior”
From Continuing challenges of elephant captivity: the captive environment, health issues, and welfare implications (2024) · cites this paper as Greco et al., 2016a
In the U.S. most captive elephants have experienced at least one inter-zoo transfer, which is associated with stereotypic behaviors.
“For example, in the U.S. most captive elephants have experienced at least one inter-zoo transfer (111), which is associated with stereotypic behaviors (35).”
From Comparison of animal welfare assessment tools and methodologies: need for an effective approach for captive elephants in Asia (2024) · cites this paper as [35]
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- Authors
- Greco, B. J., Meehan, C. L., Hogan, J. N., Leighty, K. A., Mellen, J., Mason, G. J., & Mench, J. A.
- Affiliations
- University of California, Davis (US)
- University of Guelph (CA)
- Corresponding author
- Greco, Brian J.
- Venue
- PLoS ONE
- Publisher
- Public Library of Science (PLoS)
- Published
- 2016-07-14
- Volume
- 11
- Issue
- 7
- Pages
- e0144276
- ISSN
- 1932-6203 (ISSN-L)
- PMC
- PMC4944985
- OpenAlex
- W2465848062
- Cited by
- 94
- References
- 61
- Citations by year
- 2026: 8 · 2025: 3 · 2024: 9 · 2023: 6 · 2022: 8 · 2021: 12 · 2020: 11 · 2019: 16 · 2018: 8 · 2017: 4 · 2016: 7
- FWCI
- 17.88
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Licence
- Locations
- PLoS ONE (publishedVersion, cc-by)
- Europe PMC (PubMed Central) (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion)
- Funders
- Institute of Museum and Library Services (LG-25-10-0033-10)
- Topics
- Health Sciences › Veterinary › Small Animals › Animal Behavior and Welfare Studies
- Physical Sciences › Environmental Science › Ecology › Wildlife Ecology and Conservation
- Health Sciences › Veterinary › Equine › Veterinary Equine Medical Research
- Keywords
- zoo elephants, stereotypy, animal welfare, Loxodonta africana, Asian elephant, social group size, residential environment, seasonal variation, nocturnal activity, captive elephants, animal-assisted therapy, animal-assisted education
- MeSH terms
- Age Factors, Animals, Animal Welfare, Animals, Zoo, Behavior, Animal, Circadian Rhythm, Elephants, Female, Male, Motor Activity, Seasons, Social Environment, Species Specificity, Stereotyped Behavior, United States
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