The influence of forage, protected areas, and mating prospects on grouping patterns of male elephants
Abstract
Factors affecting social group size in mammals are relatively well studied for females, but less is known about determinants of group size for males, particularly in species that live in sexually segregated groups. Male grouping patterns are thought to be driven more by spatial and temporal dispersion of mating opportunities than by food resources or predation risk. We evaluated the influence of 3 factors on male group sizes and number of males in mixed-sex groups in African elephants; forage availability (using Normalized Difference Vegetation Index, a satellite-based indicator of primary productivity), anthropogenic mortality risk (using distance of elephants from a protected area center), and mating opportunities (using the number of males in mixed-sex groups with and without estrous females). Using zero-truncated negative binomial regressions and a model-selection approach, we found that male elephants occurred in larger groups where primary productivity was higher and where they were further from a protected area center. However, we found an interaction between primary productivity and anthropogenic mortality risk: at low primary productivity, elephants formed larger groups further away from a protected area center, but did less so at higher primary productivity. This pattern suggests that male elephants are sensitive to seasonal variation in potential anthropogenic mortality risk, by remaining in smaller groups when risk is low, but forming larger groups when risk is high. Mating opportunities also led to an increase in male numbers in mixed-sex groups, but its relative influence on male grouping was less important because mating opportunities were rare.
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Male elephants in Kenya were observed in all-male groups 46% of the time.
“For example, male elephants in Kenya were observed in all-male groups 46% of the time [14]”
From Socializing a group of male Asian elephants in a semi-captive facility in Lao PDR (2025) · cites this paper as [14]
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The Behavioral Responses of Elephants to the Maasai in Amboseli
Kangwana, K. (2011) · University of Chicago Press eBooks · doi:10.7208/chicago/9780226542263.003.0020 · Cited by 6 · see citing works
20. The Behavioral Responses of Elephants to the Maasai in Amboseli
Kangwana, K. (2019) · doi:10.7208/9780226542263-022 · Cited by 4 · see citing works
The Maasai-Elephant Relationship
Browne‐Nuñez, C. (2011) · University of Chicago Press eBooks · doi:10.7208/chicago/9780226542263.003.0019 · Cited by 3 · see citing works
Browne‐Nuñez, C. (2019) · doi:10.7208/9780226542263-021 · Cited by 1 · see citing works
Flagged first, then by citations
3 of the 117 references couldn’t be matched to a known paper.
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Mortality Patterns of Asian Elephants in a Region of Human–Elephant Conflict
LaDue, C. A., Eranda, I., Jayasinghe, C., & Vandercone, R. P. G. (2021) · The Journal of Wildlife Management · doi:10.1002/jwmg.22012 · Cited by 37 · see citing works
LaDue, C. A., Schulte, B. A., Kiso, W. K., & Freeman, E. W. (2021) · Behaviour · doi:10.1163/1568539x-bja10120 · Cited by 28 · see citing works
Anoop, N. R., Krishnan, S., & Ganesh, T. (2023) · Frontiers in Conservation Science · doi:10.3389/fcosc.2023.1142325 · Cited by 28 · see citing works
DNA methylation signatures of early-life adversity are exposure-dependent in wild baboons
Anderson, J. A., Lin, D., Lea, A. J., Johnston, R. A., Voyles, T., Akinyi, M. Y., Archie, E. A., Alberts, S. C., & Tung, J. (2024) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.2309469121 · Cited by 26 · see citing works
LaDue, C. A., Vandercone, R. P. G., Kiso, W. K., & Freeman, E. W. (2022) · Animals · doi:10.3390/ani12091215 · Cited by 26 · see citing works
The Chemical Ecology of Elephants: 21st Century Additions to Our Understanding and Future Outlooks
Schulte, B. A., & LaDue, C. A. (2021) · Animals · doi:10.3390/ani11102860 · Cited by 24 · see citing works
Cohort consequences of drought and family disruption for male and female African elephants
Lee, P. C., Moss, C. J., Njiraini, N., Poole, J. H., Sayialel, K. N., & Fishlock, V. (2021) · Behavioral Ecology · doi:10.1093/beheco/arab148 · Cited by 21 · see citing works
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Elephants move faster in small fragments of low productivity in Amboseli ecosystems: Kenya
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Schüßler, D., Lee, P. C., & Stadtmann, R. (2018) · Landscape Ecology · doi:10.1007/s10980-018-0728-7 · Cited by 19 · see citing works
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Boult, V. L., Sibly, R. M., Quaife, T. L., Fishlock, V., Moss, C., & Lee, P. C. (2018) · African Journal of Ecology · doi:10.1111/aje.12553 · Cited by 13 · see citing works
du Plessis, K., Ganswindt, S. B., Bertschinger, H. J., Crossey, B., Henley, M. D., Ramahlo, M., & Ganswindt, A. (2021) · Animals · doi:10.3390/ani11113070 · Cited by 10 · see citing works
Wilson, J. W., Bergl, R. A., Minter, L. J., Loomis, M. R., & Kendall, C. J. (2019) · International Zoo Yearbook · doi:10.1111/izy.12216 · Cited by 10 · see citing works
Agent-based modelling as a tool for elephant poaching mitigation
Neil, E., Madsen, J. K., Carrella, E., Payette, N., & Bailey, R. M. (2020) · Ecological Modelling · doi:10.1016/j.ecolmodel.2020.109054 · Cited by 11 · see citing works
LaDue, C. A., Vandercone, R. P. G., Kiso, W. K., & Freeman, E. W. (2021) · Wildlife Research · doi:10.1071/wr20175 · Cited by 10 · see citing works
Edge, A. C., Henley, M. D., Daday, J. K., & Schulte, B. A. (2017) · Human Dimensions of Wildlife · doi:10.1080/10871209.2017.1298168 · Cited by 9 · see citing works
Smit, J. B., Searle, C. E., Buchanan‐Smith, H. M., Strampelli, P., Mkuburo, L., Kakengi, V. A., Kohi, E. M., Dickman, A., & Lee, P. C. (2022) · African Journal of Ecology · doi:10.1111/aje.13083 · Cited by 7 · see citing works
Worku, E. A., Bro‐Jørgensen, J., Evangelista, P., Bekele, A., Atickem, A., & Stenseth, N. C. (2023) · Global Ecology and Conservation · doi:10.1016/j.gecco.2023.e02546 · Cited by 6 · see citing works
Elephant Scar Prevalence in the Kasigau Wildlife Corridor, Kenya: Echoes of Human-Elephant Conflict
Hagen, L. V., LaDue, C. A., & Schulte, B. A. (2023) · Animals · doi:10.3390/ani13040605 · Cited by 5 · see citing works
Fruit availability and human disturbance influence forest elephant group size
Meier, A. C., Bourgeois, S., Adams, E. M., Bikang, H. B. A., Jasperse‐Sjolander, L., Lewis, M., Masseloux, J., Morin, D. J., & Poulsen, J. R. (2023) · Animal Behaviour · doi:10.1016/j.anbehav.2023.07.002 · Cited by 5 · see citing works
Readyhough, T. S., Davis, M., Joseph, S., Moresco, A., & Schreier, A. L. (2023) · Journal of Zoological and Botanical Gardens · doi:10.3390/jzbg4010018 · Cited by 4 · see citing works
Frequency and Density Associated Grouping Patterns of Male Roosevelt Elk
Weckerly, F. W. (2020) · Frontiers in Ecology and Evolution · doi:10.3389/fevo.2020.00204 · Cited by 4 · see citing works
Sutthiboriban, N., Simcharoen, A., Gale, G. A., Ngoprasert, D., Chutipong, W., & Tantipisanuh, N. (2024) · Pacific Conservation Biology · doi:10.1071/pc23061 · Cited by 3 · see citing works
Males Matter: On the Status and Future Direction of Male Elephant Management in EAZA Institutions
LaDue, C. A., & Schiffmann, C. L. (2025) · Zoo Biology · doi:10.1002/zoo.70040 · Cited by 2 · see citing works
Suttidate, N., Radeloff, V. C., & Sukmasuang, R. (2025) · Global Ecology and Conservation · doi:10.1016/j.gecco.2025.e03945 · Cited by 2 · see citing works
Distribution and abundance of African elephants in Ngorongoro Crater, northern Tanzania
Kioko, J., Horton, A., Libre, M., Vickers, J., Dressel, E., Kasey, H., Ndegeya, P. M., Gadiye, D. E., Kissui, B. M., & Kiffner, C. (2020) · African Zoology · doi:10.1080/15627020.2020.1813625 · Cited by 2 · see citing works
Socializing a group of male Asian elephants in a semi-captive facility in Lao PDR
López Pérez, A. B., Brown, J. L., Goldenberg, S. Z., Rey, A., & Khonmee, J. (2025) · PLOS One · doi:10.1371/journal.pone.0332944 · Cited by 2 · see citing works
“The amount of time that wild males spend alone or with conspecifics depends on various factors, including age and reproductive state [18,19,21,22], forage abundance, kin relationships [14], and human activities [16].”
Mramba, R. P., & Kisanga, A. C. (2025) · Discover Animals · doi:10.1007/s44338-025-00070-1 · Cited by 1 · see citing works
Elephants’ habitat use and behaviour when outside of Gonarezhou National Park
Mandinyenya, B., Mingione, M., Traill, L. W., & Attorre, F. (2025) · Koedoe · doi:10.4102/koedoe.v67i1.1842 · Cited by 1 · see citing works
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Migration patterns of African elephants (Loxodonta africana) in Ol Pejeta Conservancy
Grabbe, E. (2018) · Epsilon Archive for Student Projects (University of Southampton) · Cited by 1 · see citing works
Mramba, R. P. (2022) · Research Square · doi:10.21203/rs.3.rs-1744068/v1
Abundance and social organisation of male Asian elephants in northwestern India
Madhusudanan, A., Natarajan, L., Gogoi, K., Mondol, S., & Pandav, B. (2025) · Wildlife Research · doi:10.1071/wr25098
Pérez, A. B. L., Goldenberg, S. Z., Brown, J. L., & Khonmee, J. (2026) · Applied Animal Behaviour Science · doi:10.1016/j.applanim.2026.107067
Moss, C. J., Lee, P. C., Njiraini, N., Sayialel, C., & Fishlock, V. (2026) · Animal Behaviour · doi:10.1016/j.anbehav.2026.123733
- Authors
- Chiyo, P. I., Wilson, J. W., Archie, E. A., Lee, P. C., Moss, C. J., & Alberts, S. C.
- Affiliations
- University of Notre Dame (US)
- The Ohio State University (US)
- University of Stirling (GB)
- Duke University (US)
- ORCID
- Venue
- Behavioral Ecology
- Publisher
- Oxford University Press (OUP)
- Published
- 2014
- Volume
- 25
- Issue
- 6
- Pages
- 1494-1504
- ISSN
- 1045-2249 (ISSN-L)
- 1465-7279
- OpenAlex
- W2162456164
- Cited by
- 37
- References
- 117
- Citations by year
- 2026: 3 · 2025: 6 · 2024: 2 · 2023: 5 · 2022: 3 · 2021: 7 · 2020: 3 · 2019: 2 · 2018: 3 · 2017: 1 · 2016: 2
- FWCI
- 0.87
- Type
- journal article
- Language
- en
- Open access
- closed
- Locations
- Behavioral Ecology (publishedVersion)
- Stirling Online Research Repository (University of Stirling) (submittedVersion)
- RePEc: Research Papers in Economics (submittedVersion)
- Topics
- Physical Sciences › Environmental Science › Ecology › Wildlife Ecology and Conservation
- Life Sciences › Agricultural and Biological Sciences › Ecology, Evolution, Behavior and Systematics › Animal Behavior and Reproduction
- Social Sciences › Psychology › Social Psychology › Primate Behavior and Ecology
- Keywords
- Loxodonta africana, social group size, forage availability, primary productivity, protected areas, mating opportunities, mixed-sex groups, sexual segregation, estrous females, seasonal variation
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