Elephant management in North American zoos: environmental enrichment, feeding, exercise, and training
Open accessAbstract
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.
The claims and statistics of this paper haven’t been checked yet.
Run the check — free, no account needed. It reads the open-access text and checks the sources it cites.
Integrity signals
Registry
Cites
Statistics
Claims the paper makes
The claims and statistics of this paper haven’t been checked yet.
Run the check — free, no account needed. It reads the open-access text and checks the sources it cites.
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
Problem-solving opportunities were components in the enrichment programs of 97% of 63 zoos examined, but these opportunities were provided infrequently.
“Greco et al. (2016b) found that problem-solving opportunities were components in the enrichment programs of 97% of the 63 zoos they examined; however, these opportunities were provided infrequently.”
From Continuing challenges of elephant captivity: the captive environment, health issues, and welfare implications (2024) · cites this paper as Greco et al., 2016b
There are few studies on the effects of enrichment for elephants, with most focused on feeding enrichment.
“There are few studies on the effects of enrichment for elephants, with most of them focused on feeding enrichment (Greco et al., 2016b).”
From Continuing challenges of elephant captivity: the captive environment, health issues, and welfare implications (2024) · cites this paper as Greco et al., 2016b
Cognitive enrichment may be provided infrequently in zoos, with keepers perceiving semi-permanent and permanent exhibit features as more important than cognitive enrichment.
“cognitive enrichment may be provided infrequently in zoos, with keepers perceiving that semi-permanent and permanent exhibit features (e.g., pools, logs, scratching areas, puzzle feeders) are more important than cognitive enrichment (Hoy, Murray & Tribe, 2010;Greco et al., 2016b).”
From Continuing challenges of elephant captivity: the captive environment, health issues, and welfare implications (2024) · cites this paper as Greco et al., 2016b
The frequency of problem-solving enrichment was so low that Greco et al. (2016b) were unable to test its effects in their welfare models.
“In fact, the frequency of problem-solving enrichment was so low that Greco et al. (2016b) were unable to test its effects in their welfare models.”
From Continuing challenges of elephant captivity: the captive environment, health issues, and welfare implications (2024) · cites this paper as Greco et al., 2016b
Asian elephants are managed for a significantly greater proportion of their day than African elephants, who tend to be more independent.
“Asian elephants are managed for a significantly greater proportion of their day than African elephants, who tend to be more independent”
From Continuing challenges of elephant captivity: the captive environment, health issues, and welfare implications (2024) · cites this paper as Greco et al., 2016b
Self-check
Percentages, sums and reported differences re-computed from the paper’s own numbers.
The self-check needs the paper’s full text.
Cites
Generalized Estimating Equations
Hardin, J. W., & Hilbe, J. M. (2002) · doi:10.1201/9781420035285 · Cited by 1,766 · see citing works
Born to choose: the origins and value of the need for control
Leotti, L. A., Iyengar, S. S., & Ochsner, K. N. (2010) · Trends in Cognitive Sciences · doi:10.1016/j.tics.2010.08.001 · Cited by 717 · see citing works
Morfeld, K. A., Meehan, C. L., Hogan, J. N., & Brown, J. L. (2016) · PLOS ONE · doi:10.1371/journal.pone.0155146 · Cited by 114 · see citing works
Self-citationGreco, B. J., Meehan, C. L., Hogan, J. N., Leighty, K. A., Mellen, J., Mason, G. J., & Mench, J. A. (2016) · PLoS ONE · doi:10.1371/journal.pone.0144276 · Cited by 94 · see citing works
Self-citationBrown, J. L., Paris, S., Prado-Oviedo, N. A., Meehan, C. L., Hogan, J. N., Morfeld, K. A., & Carlstead, K. (2016) · PLoS ONE · doi:10.1371/journal.pone.0145673 · Cited by 83 · see citing works
Miller, M. A., Hogan, J. N., & Meehan, C. L. (2016) · PLoS ONE · doi:10.1371/journal.pone.0155223 · Cited by 79 · see citing works
Flagged first, then by citations
Research works printed but not linked: 42
- [1]Mason GJ, Veasey JS (2010) How should the psychological well-being of zoo elephants be objectively investigated? Zoo Biol 29: 237– 255. 10.1002/zoo.20256 19514018 · DOI
- [4]Melfi VA (2009) There are big gaps in our knowledge, and thus approach, to zoo animal welfare: a case for evidence-based zoo animal management. Zoo Biol 28: 574– 588. 10.1002/zoo.20288 19876912 · DOI · Paper page
- [6]Sukumar R (2003) The living elephants: evolutionary ecology, behavior, and conservation New York: Oxford University Press.
- [7]Prado-Oviedo NA, Bonaparte-Saller MK, Malloy E, Meehan CL, Mench JA, Carlstead K. et al (2016) Evaluation of demographics and social life events of Asian (Elephas maximus) and African elephants (Loxodonta africana) in North American zoos. PLoS One pone.0154750. PMC4945030 27415437
- [8]Wemmer C, Christen CA, editors (2008) Elephants and ethics: toward a morality of coexistence Baltimore: Johns Hopkins University Press. 483 p.
- [10]Holdgate MR, Meehan CL, Hogan JN, Miller LJ, Soltis J, Andrews J, et al (2016) Walking behavior of zoo elephants: associations between GPS-measured daily walking distances and enviornmental factors, social factors, and welfare indicators. PLoS One pone.0150331. PMC4944962 27414411
- [11]Holdgate MR, Meehan CL, Hogan JN, Miller LJ, Rushen J, de Passillé AM, et al (2016) Recumbence behavior in zoo elephants: determination of patterns and frequency of recumbent rest and associated enviornmental and social factors. PLoS One pone.0153301. PMC4945027 27414809
- [15]Meehan CL, Hogan JN, Bonaparte-Saller MK, Mench JA (2016) Housing and social enviornments of African (Loxodonta africana) and Asian (Elephas maximus) elephants in North American zoos. PLoS One pone.0146703. PMC4944928 27414034
- [16]Lambeth SP, Hau J, Perlman JE, Martino M, Schapiro SJ (2006) Positive reinforcement training affects hematologic and serum chemistry values in captive chimpanzees (Pan troglodytes). Am J Primatol 68: 245– 256. 16477594
- [21]Young RJ (2003) Environmental enrichment for captive animals Oxford: Blackwell Science Ltd. 240 p.
- [22]Shyne A (2006) Meta-analytical review of the effects of enrichment on stereotypic behavior in zoo mammals. Zoo Biol 25: 317– 337.
- [23]Swaisgood RR, Shepherdson D (2006) Environmental enrichment as a strategy for mitigating stereotypies in zoo animals: a literature review and meta-analysis In: Mason GJ, Rushen J, editors. Stereotypic animal behavior: fundamentals and applications to welfare. 2 ed: CABI.
- [24]Carlstead K, Shepherdson D (2000) Aleviating stress in zoo animals with environmental enrichment In: Moberg GP, Mench JA, editors. The biology of animal stress: basic principles and implications for animal welfare. Wallingford: Cabi Publishing pp. 337– 354.
- [25]Stoinski TS, Daniel E, Maple TL (2000) A preliminary study of the behavioral effects of feeding enrichment on African elephants. Zoo Biol 19: 485– 493. 11180410
- [28]Rees PA (2009) Activity budgets and the relationship between feeding and stereotypic behaviors in Asian elephants (Elephas maximus) in a Zoo. Zoo Biol 28: 79– 97. 10.1002/zoo.20200 19367622 · DOI
- [33]Lewis KD, Shepherdson DJ, Owens TM, Keele M (2010) A survey of elephant husbandry and foot health in North American zoos. Zoo Biol 29: 221– 236. 10.1002/zoo.20291 20014111 · DOI
- [35]Atlantis E, Barnes EH, Singh MA (2006) Efficacy of exercise for treating overweight in children and adolescents: a systematic review. Int J Obes (Lond) 30: 1027– 1040. 16534526
- [36]Henriksen EJ (2002) Invited review: Effects of accute exercise and exercise training on insulin resistance. J Appl Physiol 93: 778– 796. 12133893
- [37]Metsios GS, Stavropoulos-Kalinoglou A, Veldhuijzen van Zanten JJ, Treharne GJ, Panoulas VF, Douglas KM, et al (2008) Rheumatoid arthritis, cardiovascular disease and physical exercise: a systematic review. Rheumatology (Oxford) 47: 239– 248. 18045810
- [38]Walsmith J, Roubenoff R (2002) Cachexia in rheumatoid arthritis. Int J Cardiol 85: 89– 99. 12163213
- [39]Bell RR, Spencer MJ, Sherriff JL (1995) Diet-induced obesity in mice can be treated without energy restriction using exercise and or a low fat diet. J Nutr 125: 2356– 2363. 7666253
- [40]Honda A, Sogo N, Nagasawa S, Shimizu T, Umemuara Y (2003) High-impace exercise sterenthens bone in osteopenic ovariestomized rats with the same outcome as Sham rats. J Appl Physiol 95: 1032– 1037. 12754179
- [41]Muths E, Reichman OJ (1996) Kangaroo rat bone compared to white rat after short-term disuse and exercise. Comp Biochem Physiol 114A: 355– 361. 8759284
- [42]Rolls BJ, Rowe EA (1979) Exercise and the development and persistence of dietary obesity in male and female rats. Physiol Behav 23: 241– 247. 504410
- [43]Degeling C, Burton L, McCormack GR (2012) An investigation of the association between socio-demographic factors, dog-exercise requirements, and the amount of walking dogs receive. Can J Vet Res 76: 235– 240. 23277705 PMC3384289
- [44]Greene LM, Marcellin-Little DJ, Lascelles BDX (2013) Association among exercise duration, lameness severity, and hip joint range of motion in labrador retrievers with hip dysplasia. J Am Vet Med Assoc 242: 1528– 1533. 10.2460/javma.242.11.1528 23683017 · DOI
- [45]Linder D, Mueller M (2014) Pet obesity management: beyond nutrition. Vet Clin North Am Small Anim Pract 44: 789– 806. 10.1016/j.cvsm.2014.03.004 24951347 · DOI
- [46]Forsythe WA, Miller ER, Curry B, Bennink R (1981) Aerobic exercise effects on lipoproteins and tissue lipids in young pigs. Atherosclerosis 38: 327– 337. 7225172
- [47]Lehnhardt J, Galloway M (2008) Carrots and sticks, people and elephants: rank, domination, and training In: Wemmer C, Christen CA, editors. Elephants and ethics: toward a morality of coexistence. Baltimore: Johns Hopkins University Press pp. 167– 184.
- [50]Zeligs JA (2014) Animal training 101: the complete and practical guide to the art and science of behavior modification: Mill City Press, Inc.
- [52]Franklin JCW (1972) Punishment and frustration: Serendipitous findings. Psychol Rep: 655– 661. 5024947
- [53]Gerber B, Yarali A, Diegelmann S, Wotjak CT, Pauli P, Fendt M (2014) Pain-relief learning in flies, rats, and man: basic research and applied perspectives. Learn Mem 21: 232– 252. 10.1101/lm.032995.113 24643725 PMC3966540 · DOI
- [55]Bradshaw GA (2009) Elephants on the edge: What animals teach us about humanity New Haven, CT: Yale University Press. 352 p.
- [58]Holgado–Tello FP, Chacón–Moscoso S, Barbero–García I, Vila–Abad E (2010) Polychoric versus Pearson correlations in exploratory and confirmatory factor analysis of ordinal variables. Qual Quant 44: 153– 166.
- [62]Cohen J, Cohen P, West SG, Aiken LS (2003) Applied multiple regression/correlation analysis for the behavioral sciences New York: Routledge.
- [63]Hoy JM, Murray PJ, Tribe A (2010) Thirty years later: enrichment practices for captive mammals. Zoo Biol 29: 303– 316. 10.1002/zoo.20254 19434736 · DOI · Paper page
- [66]Bennett C, Torgerson-White L, Fripp D, Watters J, Petric A (2015) A Multi-Institutional Assessment of Factors Influencing Locomotion and Pacing in Captive Okapis (Okapia johnstoni). J Appl Anim Welf Sci 18 Suppl 1: S43– 61. 10.1080/10888705.2015.1075835 26440496 · DOI
- [67]Schneider JD, Tokach MD, Dritz SS, Nelssen JL, Derouchey JM, Goodband RD (2007) Effects of feeding schedule on body condition, aggressiveness, and reproductive failure in group-housed sows. J Anim Sci 85: 3462– 3469. 17785592
- [68]Bashaw MJ, Tarou LR, Maki TS, Maple TL (2001) A survey assessment of variables related to stereotypy in captive giraffe and okapi. Appl Anim Behav Sci 73: 235– 247. 11376840
- [71]Mistlberger RE (1994) Circadian food-anticipatory activity: formal models and physiological mechanisms. Neurosci Biobehav R 18: 171– 195. 8058212
- [72]Laule G, Desmond T (1998) Postitive reinforcement training as an enrichment strategy. In: Shepherdson D, Mellen JD, Hutchins M, editors. Second nature: Environmental enrichment for captive animals. pp. 302–313.
- [76]Butterworth A, Mench JA, Wielebnowski N (2011) Practical Strategies to Assess (and Improve) Welfare In: Appleby MC, Mench JA, Olsson IAS, Hughes BO, editors. Animal Welfare. 2 ed: CAB International pp. 200– 214.
Cited by
Sorted by citations
Morfeld, K. A., Meehan, C. L., Hogan, J. N., & Brown, J. L. (2016) · PLOS ONE · doi:10.1371/journal.pone.0155146 · Cited by 114 · see citing works
Self-citationGreco, B. J., Meehan, C. L., Hogan, J. N., Leighty, K. A., Mellen, J., Mason, G. J., & Mench, J. A. (2016) · PLoS ONE · doi:10.1371/journal.pone.0144276 · Cited by 94 · see citing works
Self-citationBrown, J. L., Paris, S., Prado-Oviedo, N. A., Meehan, C. L., Hogan, J. N., Morfeld, K. A., & Carlstead, K. (2016) · PLoS ONE · doi:10.1371/journal.pone.0145673 · Cited by 83 · see citing works
Miller, M. A., Hogan, J. N., & Meehan, C. L. (2016) · PLoS ONE · doi:10.1371/journal.pone.0155223 · Cited by 79 · see citing works
Holdgate, M. R., Meehan, C. L., Hogan, J. N., Miller, L. J., Soltis, J., Andrews, J., & Shepherdson, D. J. (2016) · PLoS ONE · doi:10.1371/journal.pone.0150331 · Cited by 65 · see citing works
Self-citationGreco, B. J., Meehan, C. L., Heinsius, J. L., & Mench, J. A. (2017) · Applied Animal Behaviour Science · doi:10.1016/j.applanim.2017.05.003 · Cited by 56 · see citing works
Self-citationBrown, J. L., Carlstead, K., Bray, J. D., Dickey, D., Farin, C., & Ange-van Heugten, K. (2019) · PLOS ONE · doi:10.1371/journal.pone.0217326 · Cited by 46 · see citing works
Self-citationVeasey, J. S. (2019) · Animals · doi:10.3390/ani10010039 · Cited by 36 · see citing works
Scott, N. L., & LaDue, C. A. (2019) · Zoo Biology · doi:10.1002/zoo.21506 · Cited by 30 · see citing works
Brown, J. L., Bansiddhi, P., Khonmee, J., & Thitaram, C. (2020) · Animals · doi:10.3390/ani10040737 · Cited by 26 · see citing works
Fazio, J. M., Barthel, T., Freeman, E. W., Garlick-Ott, K., Scholle, A., & Brown, J. L. (2020) · Animals · doi:10.3390/ani10112026 · Cited by 24 · see citing works
Morfeld, K. A., & Brown, J. L. (2017) · PLOS ONE · doi:10.1371/journal.pone.0188701 · Cited by 22 · see citing works
Self-citationDoyle, C., Rally, H., O’Brien, L., Tennison, M., Marino, L., & Jacobs, B. (2024) · PeerJ · doi:10.7717/peerj.18161 · Cited by 14 · see citing works
Innovating to solve a novel puzzle: wild Asian elephants vary in their ability to problem solve
Jacobson, S. L., Dechanupong, J., Horpiencharoen, W., Yindee, M., & Plotnik, J. M. (2023) · Animal Behaviour · doi:10.1016/j.anbehav.2023.08.019 · Cited by 14 · see citing works
Snyder, R. J., Barrett, L. P., Emory, R. A., & Perdue, B. M. (2021) · Animal Cognition · doi:10.1007/s10071-021-01504-5 · Cited by 11 · see citing works
- Authors
- Greco, B. J., Meehan, C. L., Miller, L. J., Shepherdson, D. J., Morfeld, K. A., Andrews, J., Baker, A. M., Carlstead, K., & Mench, J. A.
- Affiliations
- University of California, Davis (US)
- Brookfield Zoo Chicago (US)
- Oregon Zoo (US)
- Smithsonian Conservation Biology Institute (US)
- Corresponding author
- Greco, Brian J.
- Venue
- PLOS ONE
- Publisher
- Public Library of Science (PLoS)
- Published
- 2016-07-14
- Volume
- 11
- Issue
- 7
- Pages
- e0152490
- ISSN
- 1932-6203 (ISSN-L)
- PMC
- PMC4944993
- OpenAlex
- W2468496484
- Cited by
- 73
- References
- 6
- Citations by year
- 2026: 6 · 2025: 3 · 2024: 7 · 2023: 10 · 2022: 5 · 2021: 4 · 2020: 10 · 2019: 10 · 2018: 6 · 2017: 3 · 2016: 9
- FWCI
- 14.35
- 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)
- RePEc: Research Papers in Economics (submittedVersion)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
- 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
- elephant management, zoo elephants, Loxodonta africana, Asian elephant, environmental enrichment, feeding schedule, physical activity, animal training, animal welfare, zoo animal husbandry, Association of Zoos and Aquariums, sex differences, species differences
- MeSH terms
- Animals, Animal Husbandry, Animal Welfare, Animals, Zoo, Elephants, Environment, Feeding Behavior, Female, Housing, Animal, Male, North America, Physical Conditioning, Animal, Social Environment
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
Are you an author, or do you see a problem with this page? Request a review