Mapping shade availability and use in zoo environments: a tool for evaluating thermal comfort
Open accessAbstract
For many species in zoos, particularly megafauna vulnerable to heat stress, shade is a key environmental resource. However, shade availability has received comparatively less attention than other aspects of the zoo environment. In this study, we share a simple low-cost approach that we applied to document shade availability across 33 zoo enclosures. We then combined these assessments with behavioral observations of enclosure use and shade-seeking behavior during summer months in a case study focused on Sichuan takin (Budorcas taxicolor tibetana) (n = 3), a large cold-adapted bovid. Behavioral observations were conducted before and after installation of a shade sail for the takin. Results indicated that shade availability varied widely across zoo enclosures, with the percent of shaded space ranging from 85 % to 22 % across enclosures during summer months. Shade was a dynamic resource and increased throughout the year and fluctuated across the day, with the least shade available midday. Takin showed general preferences for shaded areas near the walls of their enclosure but were observed using newly available shade from the shade sail after its installation. These accessible methods can be easily applied to assess shade within existing enclosures, evaluate enclosure modifications, and provide guidance for the design of new enclosures.
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.
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
nlme : Linear and nonlinear mixed effects models
DebRoy, S. (2006) · Medical Entomology and Zoology · Cited by 10,009 · see citing works
nlme: Linear and Nonlinear Mixed Effects Models
Pinheiro, J., R Core Team, & Bates, D. (1999) · doi:10.32614/cran.package.nlme · Cited by 6,867 · see citing works
Impact of Heat Stress on Poultry Production
Lara, L. J. C., & Rostagno, M. H. (2013) · Animals · doi:10.3390/ani3020356 · Cited by 1,276 · see citing works
Measuring Behaviour: An Introductory Guide
Martin, P., & Bateson, P. (2007) · doi:10.1017/cbo9780511810893 · Cited by 1,008 · see citing works
A review of human thermal comfort in the built environment
Rupp, R. F., Vásquez, N. G., & Lamberts, R. (2015) · Energy and Buildings · doi:10.1016/j.enbuild.2015.07.047 · Cited by 907 · see citing works
Invited review: Effects of heat stress on dairy cattle welfare
Polsky, L., & von Keyserlingk, M. A. G. (2017) · Journal of Dairy Science · doi:10.3168/jds.2017-12651 · Cited by 874 · see citing works
Stevenson, R. D. (1985) · The American Naturalist · doi:10.1086/284423 · Cited by 511 · see citing works
Behavioral thermoregulation in mammals: a review
Terrien, J. (2010) · Frontiers in bioscience · doi:10.2741/3797 · Cited by 402 · see citing works
Tucker, C. B., Rogers, A. R., & Schütz, K. E. (2007) · Applied Animal Behaviour Science · doi:10.1016/j.applanim.2007.03.015 · Cited by 376 · see citing works
Thermal physiology of laboratory mice: Defining thermoneutrality
Gordon, C. J. (2012) · Journal of Thermal Biology · doi:10.1016/j.jtherbio.2012.08.004 · Cited by 305 · see citing works
Poultry production in hot climates
(2008) · CABI eBooks · doi:10.1079/9781845932589.0000 · Cited by 275 · see citing works
Glossary of terms for thermal physiology (Third Edition)
IUPSThermalCommission (2001) · The Japanese Journal of Physiology · Cited by 260 · see citing works
Thermophysiological models and their applications: A review
Katić, K., Li, R., & Zeiler, W. (2016) · Building and Environment · doi:10.1016/j.buildenv.2016.06.031 · Cited by 243 · see citing works
Beyond the classic thermoneutral zone: Including thermal comfort
Kingma, B. R., Frijns, A. J., Schellen, L., & van Marken Lichtenbelt, W. D. (2014) · Temperature · doi:10.4161/temp.29702 · Cited by 207 · see citing works
Fuller, A., Mitchell, D., Maloney, S. K., & Hetem, R. S. (2016) · Climate Change Responses · doi:10.1186/s40665-016-0024-1 · Cited by 207 · see citing works
Temperature constraints on foraging behaviour of male Alpine ibex (Capra ibex) in summer
Aublet, J.-F., Festa‐Bianchet, M., Bergero, D., & Bassano, B. (2008) · Oecologia · doi:10.1007/s00442-008-1198-4 · Cited by 178 · see citing works
Translating Animal Model Research: Does It Matter That Our Rodents Are Cold?
Maloney, S. K., Fuller, A., Mitchell, D., Gordon, C. J., & Overton, J. M. (2014) · Physiology · doi:10.1152/physiol.00029.2014 · Cited by 166 · see citing works
Schütz, K. E., Rogers, A. R., Cox, N. R., & Tucker, C. B. (2008) · Applied Animal Behaviour Science · doi:10.1016/j.applanim.2008.07.005 · Cited by 152 · see citing works
Schütz, K. E., Cox, N. R., & Matthews, L. R. (2008) · Applied Animal Behaviour Science · doi:10.1016/j.applanim.2008.04.001 · Cited by 126 · see citing works
Owen‐Smith, N., & Goodall, V. (2014) · Journal of Zoology · doi:10.1111/jzo.12132 · Cited by 126 · see citing works
Causes and consequences of intraspecific variation in animal responses to anthropogenic noise
Harding, H. R., Lamont, T. A. C., Eastcott, E., Simpson, S. D., & Radford, A. N. (2019) · Behavioral Ecology · doi:10.1093/beheco/arz114 · Cited by 116 · see citing works
Thermal constraints on activity scheduling and habitat choice in baboons
Hill, R. A. (2005) · American Journal of Physical Anthropology · doi:10.1002/ajpa.20264 · Cited by 115 · see citing works
Effects of body size on the diurnal activity budgets of African browsing ruminants
Toit, D., & Yetman, C. A. (2004) · Oecologia · doi:10.1007/s00442-004-1789-7 · Cited by 101 · see citing works
Pruetz, J. D. (2007) · Primates · doi:10.1007/s10329-007-0038-1 · Cited by 100 · see citing works
Coimbra, P. A. D., Filho, L. C. P. M., & Hötzel, M. J. (2012) · Applied Animal Behaviour Science · doi:10.1016/j.applanim.2012.04.009 · Cited by 91 · see citing works
Veldhuis, M. P., Kihwele, E. S., Cromsigt, J. P. G. M., Ogutu, J. O., Hopcraft, J. G. C., Owen‐Smith, N., & Olff, H. (2019) · Ecology Letters · doi:10.1111/ele.13350 · Cited by 86 · see citing works
Monitoring the behavior and habitat use of animals to enhance welfare using the ZooMonitor app
Wark, J. D., Cronin, K. A., Niemann, T., Shender, M. A., Horrigan, A., Kao, A., & Ross, M. R. (2019) · Animal Behavior and Cognition · doi:10.26451/abc.06.03.01.2019 · Cited by 86 · see citing works
Self-citationMøller, E. F., Kamau, J. M. Z., & Maloiy, G. M. O. (1983) · Comparative Biochemistry and Physiology Part A Physiology · doi:10.1016/0300-9629(83)90608-4 · Cited by 78 · see citing works
A comparison of nocturnal primate behavior in exhibits illuminated with red and blue light
Fuller, G. A., Raghanti, M. A., Dennis, P. M., Kuhar, C. W., Willis, M. A., Schook, M. W., & Lukas, K. E. (2016) · Applied Animal Behaviour Science · doi:10.1016/j.applanim.2016.08.011 · Cited by 72 · see citing works
Chiew, S. J., Butler, K. L., Sherwen, S. L., Coleman, G. J., Fanson, K. V., & Hemsworth, P. H. (2019) · Animals · doi:10.3390/ani9060285 · Cited by 61 · see citing works
Orban, D. A., Soltis, J. M., Perkins, L., & Mellen, J. D. (2017) · Zoo Biology · doi:10.1002/zoo.21366 · Cited by 57 · see citing works
SCALING ENERGETICS OF HOMEOTHERMIC VERTEBRATES: AN OPERATIONAL ALLOMETRY
Calder, W. A. (1987) · Annual Review of Physiology · doi:10.1146/annurev.ph.49.030187.000543 · Cited by 54 · see citing works
Shrestha, A. K., van Wieren, S. E., van Langevelde, F., Fuller, A., Hetem, R. S., Meyer, L. C. R., de Bie, S., & Prins, H. H. T. (2013) · International Journal of Biometeorology · doi:10.1007/s00484-012-0622-y · Cited by 50 · see citing works
Reassessing the relationship between juvenile mortality and genetic monomorphism in captive cheetahs
Wielebnowski, N. C. (1996) · Zoo Biology · doi:10.1002/(sici)1098-2361(1996)15:4 · Cited by 45 · see citing works
A Comparison of Zoo Animal Behavior in the Presence of Familiar and Unfamiliar People
Martin, R. A., & Melfi, V. A. (2016) · Journal of Applied Animal Welfare Science · doi:10.1080/10888705.2015.1129907 · Cited by 42 · see citing works
Zeng, Z., Beck, P. S. A., Wang, T., Skidmore, A. K., Song, Y., Gong, H., & Prins, H. H. T. (2010) · Journal of Mammalogy · doi:10.1644/08-mamm-a-390r.1 · Cited by 41 · see citing works
Nielsen, B. L. (2018) · Applied Animal Behaviour Science · doi:10.1016/j.applanim.2018.04.013 · Cited by 39 · see citing works
Required thermal thresholds during transport of animals
Schrama, J. W., van der Her, W., Gorssen, J., Henken, A. M., Verstegen, M. W. A., & Noordhuizen, J. P. T. M. (1996) · Veterinary Quarterly · doi:10.1080/01652176.1996.9694624 · Cited by 35 · see citing works
Zeller, U., & Göttert, T. (2019) · Global Ecology and Conservation · doi:10.1016/j.gecco.2019.e00756 · Cited by 31 · see citing works
Powell, D., Speeg, B., Li, S., Blumer, E. S., & McShea, W. J. (2013) · Mammalia · doi:10.1515/mammalia-2012-0076 · Cited by 25 · see citing works
Woolway, E. E., & Goodenough, A. E. (2017) · Zoo Biology · doi:10.1002/zoo.21357 · Cited by 25 · see citing works
Gu, Z., Yang, S., Leng, J., Xu, S., Tang, S., Liu, C., Gao, Y., & Mao, H. (2016) · Applied Animal Behaviour Science · doi:10.1016/j.applanim.2016.01.024 · Cited by 22 · see citing works
Behavioral thermoregulation in a group of zoo‐housed colobus monkeys ( Colobus guereza )
Wark, J. D., Kuhar, C. W., & Lukas, K. E. (2014) · Zoo Biology · doi:10.1002/zoo.21138 · Cited by 9 · see citing works
Self-citationQuantifying shade using a standard environment
Langman, V. A., Rowe, M. F., Forthman, D. L., Langman, N., Black, J. D., & Walker, T. (2003) · Zoo Biology · doi:10.1002/zoo.10067 · Cited by 9 · see citing works
Flagged first, then by citations
1 of the 45 references couldn’t be matched to a known paper.
Research works printed but not linked: 1
- [24]Fanger P.O. Thermal Comfort: Analysis and Applications in Environmental Engineering Danish Technical Press Copenhagen, Denmark 1970
Cited by
Sorted by citations
de Azevedo, C. S., Cipreste, C. F., Pizzutto, C. S., & Young, R. J. (2023) · Animals · doi:10.3390/ani13081277 · Cited by 82 · 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
Wark, J. D., Wierzal, N. K., & Cronin, K. A. (2021) · Journal of Zoological and Botanical Gardens · doi:10.3390/jzbg2020014 · Cited by 24 · see citing works
Self-citationRose, P. E., & Lewton, J. (2024) · Journal of Applied Animal Welfare Science · doi:10.1080/10888705.2024.2440891 · Cited by 23 · see citing works
Do zoo‐housed primates retreat from crowds? A simple study of five primate species
Cairo-Evans, A., Wierzal, N. K., Wark, J. D., & Cronin, K. A. (2022) · American Journal of Primatology · doi:10.1002/ajp.23386 · Cited by 19 · see citing works
Self-citationArtificial Intelligence for Climate Change Biology: From Data Collection to Predictions
Levy, O., & Shahar, S. (2024) · Integrative and Comparative Biology · doi:10.1093/icb/icae127 · Cited by 19 · see citing works
Krebs, B. L., Chudeau, K. R., Eschmann, C. L., Tu, C. W., Pacheco, E., & Watters, J. V. (2022) · Frontiers in Veterinary Science · doi:10.3389/fvets.2022.972217 · Cited by 17 · see citing works
An evaluation of thermal imaging as a welfare monitoring tool for captive chimpanzees
Ross, S. R., Lake, B. R., Fultz, A. L., & Hopper, L. M. (2021) · Primates · doi:10.1007/s10329-021-00943-5 · Cited by 14 · see citing works
The Welfare of Domestic Goats (Capra hircus) in a Zoo-Based Animal-Visitor Interaction Program
Ramont, M., Leahy, M., & Cronin, K. A. (2021) · Animal Behavior and Cognition · doi:10.26451/abc.08.04.04.2021 · Cited by 13 · see citing works
Self-citationWark, J. D., Schook, M. W., Dennis, P. M., & Lukas, K. E. (2022) · American Journal of Primatology · doi:10.1002/ajp.23421 · Cited by 12 · see citing works
Self-citationPostoccupancy evaluation of staff, visitors, and three species of animals in a zoo setting
Snyder, R. J., & Barrett, L. P. (2022) · Zoo Biology · doi:10.1002/zoo.21720 · Cited by 10 · see citing works
Estimating animal location from non-overhead camera views
Woods, J. M., & Adcock, S. (2023) · Applied Animal Behaviour Science · doi:10.1016/j.applanim.2023.106032 · Cited by 10 · see citing works
Ghimire, R., Brown, J. L., Thitaram, C., & Bansiddhi, P. (2024) · Frontiers in Veterinary Science · doi:10.3389/fvets.2024.1370909 · Cited by 8 · see citing works
“ZooMonitor was developed by the Lincoln Park Zoo as a simple, software-based online tool to record the behavior and space utilization of individual animals using a digital device (72, 73).”
MethodLeveraging camera traps and artificial intelligence to explore thermoregulation behaviour
Shermeister, B., Mor, D., & Levy, O. (2024) · Journal of Animal Ecology · doi:10.1111/1365-2656.14139 · Cited by 7 · see citing works
Agrivoltaic systems for sustainability: An overview of emerging trends and practices
Maity, R., Hariram, N. P., Quazi, M. M., & Sudhakar, K. (2025) · Solar Compass · doi:10.1016/j.solcom.2025.100148 · Cited by 6 · see citing works
D’Cruze, N. C., & Groves, G. (2024) · CABI eBooks · doi:10.1079/9781800622715.0003 · Cited by 5 · see citing works
An enclosure quality ranking framework for terrestrial animals in captivity
Flanagan, A. M., Masuda, B. M., Grabar, K., Barrett, L. P., & Swaisgood, R. R. (2024) · Applied Animal Behaviour Science · doi:10.1016/j.applanim.2024.106378 · Cited by 5 · see citing works
McConnell, H., Brereton, J. E., Rice, T., & Rose, P. E. (2022) · Journal of Zoological and Botanical Gardens · doi:10.3390/jzbg3010007 · Cited by 5 · see citing works
O’Brien, S. L., Koenig, S., & Cronin, K. A. (2024) · Zoo Biology · doi:10.1002/zoo.21856 · Cited by 4 · see citing works
Self-citationIncorporating the Enclosure MonitoR Use (EMU) App to ZooMonitor Observations Technical Report
Brereton, J. E., Tuke, J., & Fernandez, E. J. (2025) · Zoo Biology · doi:10.1002/zoo.21909 · Cited by 3 · see citing works
Metrione, L. C., Pham, N. K., Price, C. A., Duskin, L., Stamper, A. M., & Penfold, L. M. (2024) · Zoo Biology · doi:10.1002/zoo.21844 · Cited by 2 · see citing works
Fitts, J., Kubasiewicz, L. M., Norris, S. L., Worth, S., Watson, T. L., Angell, R. L., Steer, M. D., & Lintott, P. R. (2025) · Animals · doi:10.3390/ani15233421 · Cited by 1 · see citing works
Organisaatioviestintä osaamisen kehittäjänä terveydenhuoltoalalla
Ottela, E. (2011) · Tampere University Institutional Repository (Tampere University)
Quantifying Use of Elevated Surfaces for Climbing Sichuan Takins (Budorcas taxicolor) in Human Care
Baskir, E. A., Callagy, C., Poelker, C., & Powell, D. (2025) · Zoo Biology · doi:10.1002/zoo.70006
Pujar, B. B., Seetharaman, K., Kumar, P., Kumar, C., Singh, A., Gnanaolivu, S. D., & Gupta, B. K. (2026) · Research Square · doi:10.21203/rs.3.rs-10882389/v1
- Authors
- Wark, J. D., Wierzal, N. K., & Cronin, K. A.
- Affiliations
- Lincoln Park Zoo (US)
- Corresponding author
- Wark, Jason D.
- Venue
- Animals
- Publisher
- MDPI AG
- Published
- 2020-07-14
- Volume
- 10
- Issue
- 7
- Pages
- 1189
- ISSN
- 2076-2615 (ISSN-L)
- PMC
- PMC7401555
- OpenAlex
- W3042685226
- Cited by
- 25
- References
- 45
- Citations by year
- 2026: 1 · 2025: 4 · 2024: 8 · 2023: 2 · 2022: 5 · 2021: 3 · 2020: 1
- FWCI
- 2.63
- Type
- journal article
- Language
- en
- Open access
- gold
- dx.doi.org
- Licence
- Locations
- Animals (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
- Topics
- Health Sciences › Veterinary › Small Animals › Animal Behavior and Welfare Studies
- Life Sciences › Agricultural and Biological Sciences › Animal Science and Zoology › Effects of Environmental Stressors on Livestock
- Physical Sciences › Environmental Science › Ecology › Physiological and biochemical adaptations
- Keywords
- zoo enclosures, thermal comfort, heat stress, enclosure design, megafauna, wildlife welfare
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