Register      Login
Wildlife Research Wildlife Research Society
Ecology, management and conservation in natural and modified habitats
RESEARCH ARTICLE

Scars of human–elephant conflict: patterns inferred from field observations of Asian elephants in Sri Lanka

Chase A. LaDue https://orcid.org/0000-0002-3224-0397 A E , Rajnish P. G. Vandercone https://orcid.org/0000-0003-1800-4133 B , Wendy K. Kiso https://orcid.org/0000-0002-6675-5970 C and Elizabeth W. Freeman https://orcid.org/0000-0001-7374-1502 D
+ Author Affiliations
- Author Affiliations

A Department of Environmental Science and Policy, George Mason University, 4400 University Drive, MS 5F2, Fairfax, VA 22030, USA.

B Department of Biological Sciences, Faculty of Applied Sciences, Rajarata University of Sri Lanka, Mihintale 50300, Sri Lanka.

C White Oak Conservation Foundation, 581705 White Oak Road, Yulee, FL 32097, USA.

D School of Integrative Studies, George Mason University, 4400 University Drive, MS 5D3, Fairfax, VA 22030, USA.

E Corresponding author. Email: cladue@gmu.edu

Wildlife Research - https://doi.org/10.1071/WR20175
Submitted: 13 October 2020  Accepted: 14 February 2021   Published online: 11 May 2021

Abstract

Context: Human–elephant conflict (HEC) is a major threat to Asian elephants as humans and elephants are forced to share common resources. In Sri Lanka, human-dominated landscapes adjacent to protected areas promote high rates of HEC, especially in the form of crop-foraging by elephants. Crop-foraging can be dangerous to both elephants and humans involved in the conflict. Gunfire is a common way for human communities to deter crop-foraging elephants, and gunshot wounds are commonly described in this elephant population on necropsy.

Aims: We sought to quantify and describe unique scar patterns among Asian elephants in a protected area, Wasgamuwa National Park, attributed to HEC.

Methods: We identified 38 adult female and 64 adult male elephants and recorded the age class and body condition of each with established standards. Using photographs, we counted the number, position, and relative size of all scars on each animal.

Key results: Male elephants had significantly more scars than did females, and for males, the number of scars increased progressively with age. Additionally, male elephants with higher body conditions had more scars. Finally, males tended to have more scars towards the head, especially at older ages.

Conclusions: Differences in total scar counts between the sexes in this population imply that male elephants in this area more frequently engage in HEC than do females, following observations previously described in the literature. Furthermore, the fact that male elephants acquired progressively more scars as they aged, and that fatter elephants had more scars, indicates that previous exposure to HEC may not have been a deterrent for future events among these males, and potentially, crops served as valuable food sources for these animals. Finally, the changing body locations of these scars with age in males possibly shows plastic behavioural responses during crop-foraging or lower tolerance by farmers towards habitual crop foragers.

Implications: These results emphasise the need for animal-based approaches to HEC mitigation. Similarly, conservation managers in Sri Lanka and other elephant range countries should investigate similar methods that estimate patterns of HEC to develop effective management strategies directly targeting animals most likely to engage in conflict.

Keywords: Asian elephant, body condition, crop-foraging, gunshot, human–elephant conflict, Sri Lanka.


References

Ahlering, M. A., Millspaugh, J. J., Woods, R. J., Western, D., and Eggert, L. S. (2011). Elevated levels of stress hormones in crop-raiding male elephants. Animal Conservation 14, 124–130.
Elevated levels of stress hormones in crop-raiding male elephants.Crossref | GoogleScholarGoogle Scholar |

Anuradha, J. M. P. N., Fujimura, M., Inaoka, T., and Sakai, N. (2019). The role of agricultural land use pattern dynamics on elephant habitat depletion and human–elephant conflict in Sri Lanka. Sustainability 11, 2818.
The role of agricultural land use pattern dynamics on elephant habitat depletion and human–elephant conflict in Sri Lanka.Crossref | GoogleScholarGoogle Scholar |

Arivazhagan, C., and Sukumar, R. (2008). Constructing age structures of Asian elephant populations: a comparison of two field methods of age estimation. Gajah 29, 11–16.

AsERSM (2017). Asian Elephant Range States Meeting final report. Ministry of Environment and Forestry, Government of Indonesia, Jakarta, Indonesia.

Bandara, R., and Tisdell, C. (2003). Comparison of rural and urban attitudes to the conservation of Asian elephants in Sri Lanka: empirical evidence. Biological Conservation 110, 327–342.
Comparison of rural and urban attitudes to the conservation of Asian elephants in Sri Lanka: empirical evidence.Crossref | GoogleScholarGoogle Scholar |

Barrett, L. P., Stanton, L. A., and Benson-Amram, S. (2019). The cognition of ‘nuisance’ species. Animal Behaviour 147, 167–177.
The cognition of ‘nuisance’ species.Crossref | GoogleScholarGoogle Scholar |

Barua, M., Bhagwat, S. A., and Jadhav, S. (2013). The hidden dimensions of human-wildlife conflict: health impacts, opportunity and transaction costs. Biological Conservation 157, 309–316.
The hidden dimensions of human-wildlife conflict: health impacts, opportunity and transaction costs.Crossref | GoogleScholarGoogle Scholar |

Bates, L. A., Poole, J. H., and Byrne, R. W. (2008). Elephant cognition. Current Biology 18, R544–R546.
Elephant cognition.Crossref | GoogleScholarGoogle Scholar | 18606119PubMed |

Blake, S., and Hedges, S. (2004). Sinking the flagship: the case of forest elephants in Asia and Africa. Conservation Biology 18, 1191–1202.
Sinking the flagship: the case of forest elephants in Asia and Africa.Crossref | GoogleScholarGoogle Scholar |

Burkholder, W. J. (2000). Use of body condition scores in clinical assessment of the provision of optimal nutrition. Journal of the American Veterinary Medical Association 217, 650–654.
Use of body condition scores in clinical assessment of the provision of optimal nutrition.Crossref | GoogleScholarGoogle Scholar | 10976293PubMed |

Campos-Arceiz, A., and Blake, S. (2011). Megagardeners of the forest: the role of elephants in seed dispersal. Acta Oecologica 37, 542–553.
Megagardeners of the forest: the role of elephants in seed dispersal.Crossref | GoogleScholarGoogle Scholar |

Campos-Arceiz, A., Takatsuki, S., Ekanayaka, S. K. K., and Hasegawa, T. (2009). The human–elephant conflict in southeastern Sri Lanka: type of damage, seasonal patterns, and sexual differences in the raiding behavior of elephants. Gajah 31, 5–14.

Chandrasekharan, K., Radhakrishnan, K., Cheeran, J., Nair, K. M., and Prabhakaran, T. (1992). Some observations on musth in captive elephants in Kerala (India). In ‘The Asian elephant: ecology, biology, disease, conservation and management’. (Eds E. Silas, M. Nair and G. Nirmalan.) pp. 71–74. (Kerala Agricultural University: Thrissur City, India.)

Chelliah, K., and Sukumar, R. (2015). Interplay of male traits, male mating strategies and female mate choice in the Asian elephant, Elephas maximus. Behaviour 152, 1113–1144.
Interplay of male traits, male mating strategies and female mate choice in the Asian elephant, Elephas maximus.Crossref | GoogleScholarGoogle Scholar |

Chiyo, P. I., Cochrane, E. P., Naughton, L., and Basuta, G. I. (2005). Temporal patterns of crop raiding by elephants: a response to changes in forage quality or crop availability? African Journal of Ecology 43, 48–55.
Temporal patterns of crop raiding by elephants: a response to changes in forage quality or crop availability?Crossref | GoogleScholarGoogle Scholar |

Chiyo, P. I., Lee, P. C., Moss, C. J., Archie, E. A., Hollister-Smith, J. A., and Alberts, S. C. (2011). No risk, no gain: effects of crop raiding and genetic diversity on body size in male elephants. Behavioral Ecology 22, 552–558.
No risk, no gain: effects of crop raiding and genetic diversity on body size in male elephants.Crossref | GoogleScholarGoogle Scholar |

Chiyo, P. I., Wilson, J. W., Archie, E. A., Lee, P. C., Moss, C. J., and Alberts, S. C. (2014). The influence of forage, protected areas, and mating prospects on grouping patterns of male elephants. Behavioral Ecology 25, 1494–1504.
The influence of forage, protected areas, and mating prospects on grouping patterns of male elephants.Crossref | GoogleScholarGoogle Scholar |

Choudhury, A., Choudhury, D. K. L., Desai, A., Duckworth, J. W., Easa, P. S., Johnsingh, A. J. T., Fernando, P., Hedges, S., Gunawardena, M., Kurt, F., Karanth, U., Lister, A., Menon, V., Riddle, H., Rübel, A., and Wikramanayake, E. (2008). Elephas maximus. In ‘The IUCN Red List of Threatened Species’. (IUCN.)

Crawley, J. A. H., Mumby, H. S., Chapman, S. N., Lahdenperä, M., Mar, K. U., Htut, W., Soe, A. T., Aung, H. H., and Lummaa, V. (2017). Is bigger better? The relationship between size and reproduction in female Asian elephants. Journal of Evolutionary Biology , .
Is bigger better? The relationship between size and reproduction in female Asian elephants.Crossref | GoogleScholarGoogle Scholar |

de Silva, S., and Wittemyer, G. (2012). A comparison of social organization in Asian elephants and African savannah elephants. International Journal of Primatology 33, 1125–1141.
A comparison of social organization in Asian elephants and African savannah elephants.Crossref | GoogleScholarGoogle Scholar |

de Silva, S., Ranjeewa, A. D. G., and Kryazhimskiy, S. (2011). The dynamics of social networks among female Asian elephants. BMC Ecology 11, 17.
The dynamics of social networks among female Asian elephants.Crossref | GoogleScholarGoogle Scholar | 21794147PubMed |

Dickerman, R. D., Pernikoff, D., Zachariah, N. Y., McConathy, W. J., Gracy, R. W., and Raven, P. V. (1994). Creatinine kinase and lactic dehydrogenase isozyme measurements in male Asian elephants (Elephas maximus) during musth and nonmusth. Clinical Chemistry 40, 989.

Edirisinghe, P. A. S., and Kitulwatte, I. G. D. (2010). Homicidal firearm injuries: a study from Sri Lanka. Forensic Science, Medicine, and Pathology 6, 93–98.
Homicidal firearm injuries: a study from Sri Lanka.Crossref | GoogleScholarGoogle Scholar |

Ekanayaka, S. K. K., Campos-Arceiz, A., Rupasinghe, M., Pastorini, J., and Fernando, P. (2011). Patterns of crop raiding by Asian elephants in a human-dominated landscape in southeastern Sri Lanka. Gajah 34, 20–25.

Evans, K. E., and Harris, S. (2008). Adolescence in male African elephants, Loxodonta africana, and the importance of sociality. Animal Behaviour 76, 779–787.
Adolescence in male African elephants, Loxodonta africana, and the importance of sociality.Crossref | GoogleScholarGoogle Scholar |

Evans, K., and Harris, S. (2012). Sex differences in habitat use by African elephants (Loxodonta africana) in the Okavango Delta, Botswana: is size really the deciding factor? African Journal of Ecology 50, 277–284.
Sex differences in habitat use by African elephants (Loxodonta africana) in the Okavango Delta, Botswana: is size really the deciding factor?Crossref | GoogleScholarGoogle Scholar |

Fernando, P. (2015). Managing elephants in Sri Lanka: where we are and where we need to be. Ceylon Journal of Science (Biological Sciences) 44, 1–11.
Managing elephants in Sri Lanka: where we are and where we need to be.Crossref | GoogleScholarGoogle Scholar |

Fernando, P., Wikramanayake, E., Weerakoon, D., Jayasinghe, L. K. A., Gunawardene, M., and Janaka, H. K. (2005). Perceptions and patterns of human-elephant conflict in old and new settlements in Sri Lanka: insights for mitigation and management. Biodiversity and Conservation 14, 2465–2481.
Perceptions and patterns of human-elephant conflict in old and new settlements in Sri Lanka: insights for mitigation and management.Crossref | GoogleScholarGoogle Scholar |

Fernando, P., Kumar, M. A., Williams, A. C., Wikramanayake, E., Aziz, T., and Singh, S. M. (2008). ‘Review of human–elephant conflict mitigation measures practiced in South Asia.’ World Bank-WWF Alliance for Forest Conservation & Sustainable Use. (World Wide Fund for Nature.)

Fernando, P., Jayewardene, J., Prasad, T., Hendavitharana, W., and Pastorini, J. (2011). Current status of Asian elephants in Sri Lanka. Gajah 35, 93–103.

Fernando, P., Leimgruber, P., Prasad, T., and Pastorini, J. (2012). Problem-elephant translocation: translocating the problem and the elephant? PLoS One 7, e50917.
Problem-elephant translocation: translocating the problem and the elephant?Crossref | GoogleScholarGoogle Scholar | 23236404PubMed |

Fernando, P., Silva, M. K. C. R. D., Jayasinghe, L. K. A., Janaka, H. K., and Pastorini, J. (2021). First country-wide survey of the endangered Asian elephant: towards better conservation and management in Sri Lanka. Oryx 55, 46–55.
First country-wide survey of the endangered Asian elephant: towards better conservation and management in Sri Lanka.Crossref | GoogleScholarGoogle Scholar |

Foerder, P., Galloway, M., Barthel, T., Moore, D. E., and Reiss, D. (2011). Insightful problem solving in an Asian elephant. PLoS One 6, e23251.
Insightful problem solving in an Asian elephant.Crossref | GoogleScholarGoogle Scholar | 21876741PubMed |

Freeman, E. W., Whyte, I., and Brown, J. L. (2009). Reproductive evaluation of elephants culled in Kruger National Park, South Africa between 1975 and 1995. African Journal of Ecology 47, 192–201.
Reproductive evaluation of elephants culled in Kruger National Park, South Africa between 1975 and 1995.Crossref | GoogleScholarGoogle Scholar |

Freeman, E. W., Schulte, B. A., and Brown, J. L. (2010). Investigating the impact of rank and ovarian activity on the social behavior of captive female African elephants. Zoo Biology 29, 154–167.
Investigating the impact of rank and ovarian activity on the social behavior of captive female African elephants.Crossref | GoogleScholarGoogle Scholar | 20391466PubMed |

Goodyear, S. E., and Schulte, B. A. (2015). Habituation to auditory stimuli by captive African elephants (Loxodonta africana). Animal Behavior and Cognition 2, 292–312.
Habituation to auditory stimuli by captive African elephants (Loxodonta africana).Crossref | GoogleScholarGoogle Scholar |

Green, M. J. B., De Alwis, S. M. D. A. U., Dayawansa, P. N., How, R., Singhakumara, B. M. P., Weerakoon, D., Wijesinghe, M. R., and Yapa, W. P. (2007). ‘Biodiversity baseline survey: Wasgomuwa National Park.’ Department of Wildlife Conservation, Ministry of Environment and Natural Resources. (ARD Inc. in association with Infotechs IDEAS and Greentech Consultants: Colombo, Sri Lanka.)

Gunawardhana, L. M. A. P. (2018). An analysis of human-elephant conflict as a disaster: a case study in Anuradhapura district of Sri Lanka. Journal of Tropical Environment 1, 40–55.

Hendavirtharana, W., Dissanayake, S., Silva, M. d., and Santiapillai, C. (1994). The survey of elephants in Sri Lanka. Gajah 12, 1–30.

Hoare, R. E. (1999). Determinants of human−elephant conflict in a land-use mosaic. Journal of Applied Ecology 36, 689–700.
Determinants of human−elephant conflict in a land-use mosaic.Crossref | GoogleScholarGoogle Scholar |

Hollister-Smith, J. A., Poole, J. H., Archie, E. A., Vance, E. A., Georgiadis, N. J., Moss, C. J., and Alberts, S. C. (2007). Age, musth and paternity success in wild male African elephants, Loxodonta africana. Animal Behaviour 74, 287–296.
Age, musth and paternity success in wild male African elephants, Loxodonta africana.Crossref | GoogleScholarGoogle Scholar |

Jadhav, S., and Barua, M. (2012). The elephant vanishes: impact of human–elephant conflict on people’s wellbeing. Health & Place 18, 1356–1365.
The elephant vanishes: impact of human–elephant conflict on people’s wellbeing.Crossref | GoogleScholarGoogle Scholar |

Jainudeen, M. R., Katongole, C. B., and Short, R. V. (1972). Plasma testosterone levels in relation to musth and sexual activity in the male Asiatic elephant, Elephas maximus. Journal of Reproduction and Fertility 29, 99–103.
Plasma testosterone levels in relation to musth and sexual activity in the male Asiatic elephant, Elephas maximus.Crossref | GoogleScholarGoogle Scholar | 5062848PubMed |

Kagwa, S. K. (2011) ‘Spatial distribution of human elephant conflict (HEC) and characterization of crop-raiding elephants in Kasigau Region, Kenya.’ (Western Kentucky University: Bowling Green, KY, USA.)

Kansky, R., and Knight, A. T. (2014). Key factors driving attitudes towards large mammals in conflict. Biological Conservation 179, 93–105.
Key factors driving attitudes towards large mammals in conflict.Crossref | GoogleScholarGoogle Scholar |

Kansky, R., Kidd, M., and Knight, A. T. (2016). A wildlife tolerance model and case study for understanding human wildlife conflicts. Biological Conservation 201, 137–145.
A wildlife tolerance model and case study for understanding human wildlife conflicts.Crossref | GoogleScholarGoogle Scholar |

Keerthipriya, P., Nandini, S., Gautam, H., Revathe, T., and Vidya, T. N. C. (2020). Musth and its effects on male–male and male–female associations in Asian elephants. Journal of Mammalogy 101, 259–270.
Musth and its effects on male–male and male–female associations in Asian elephants.Crossref | GoogleScholarGoogle Scholar |

King, L. E., Lala, F., Nzumu, H., Mwambingu, E., and Douglas-Hamilton, I. (2017). Beehive fences as multidimensional conflict-mitigation tool for farmers coexisting with elephants. Conservation Biology , .
Beehive fences as multidimensional conflict-mitigation tool for farmers coexisting with elephants.Crossref | GoogleScholarGoogle Scholar | 28221699PubMed |

Kioko, J., Muruthi, P., Omondi, P., and Chiyo, P. I. (2008). The performance of electric fences as elephant barriers in Amboseli, Kenya. South African Journal of Wildlife Research 38, 52–58.
The performance of electric fences as elephant barriers in Amboseli, Kenya.Crossref | GoogleScholarGoogle Scholar |

Kurt, F., Hartl, G. B., and Tiedemann, R. (1995). Tuskless bulls in Asian elephant Elephas maximus. History and population genetics of a man-made phenomenon. Acta Theriologica 40, 125–143.
Tuskless bulls in Asian elephant Elephas maximus. History and population genetics of a man-made phenomenon.Crossref | GoogleScholarGoogle Scholar |

LaDue, C. A., Eranda, I., Jayasinghe, C., and Vandercone, R. P. G. (2021). Mortality patterns of Asian elephants in a region of human–elephant conflict. The Journal of Wildlife Management 85, 794–802.
Mortality patterns of Asian elephants in a region of human–elephant conflict.Crossref | GoogleScholarGoogle Scholar |

Le Bel, S. (2015). Repelling elephants with a chilli pepper gas dispenser: field tests and practical use in Mozambique, Zambia and Zimbabwe. Pachyderm 56, 87–96.

Lee, P. C., and Moss, C. J. (1986). Early maternal investment in male and female African elephant calves. Behavioral Ecology and Sociobiology 18, 353–361.
Early maternal investment in male and female African elephant calves.Crossref | GoogleScholarGoogle Scholar |

Lee, P. C., and Moss, C. J. (2012). Wild female African elephants (Loxodonta africana) exhibit personality traits of leadership and social integration. Journal of Comparative Psychology 126, 224–232.
Wild female African elephants (Loxodonta africana) exhibit personality traits of leadership and social integration.Crossref | GoogleScholarGoogle Scholar | 22905995PubMed |

Leimgruber, P., Gagnon, J. B., Wemmer, C., Kelly, D. S., Songer, M. A., and Selig, E. R. (2003). Fragmentation of Asia’s remaining wildlands: implications for Asian elephant conservation. Animal Conservation 6, 347–359.
Fragmentation of Asia’s remaining wildlands: implications for Asian elephant conservation.Crossref | GoogleScholarGoogle Scholar |

Luck, C. P., and Wright, P. G. (1964). Aspects of the anatomy and physiology of the skin of the hippopotamus (H. amphibius). Quarterly Journal of Experimental Physiology and Cognate Medical Sciences 49, 1–14.
Aspects of the anatomy and physiology of the skin of the hippopotamus (H. amphibius).Crossref | GoogleScholarGoogle Scholar | 14115273PubMed |

Lynsdale, C. L., Mumby, H. S., Hayward, A. D., Mar, K. U., and Lummaa, V. (2017). Parasite-associated mortality in a long-lived mammal: variation with host age, sex, and reproduction. Ecology and Evolution 7, 10904–10915.
Parasite-associated mortality in a long-lived mammal: variation with host age, sex, and reproduction.Crossref | GoogleScholarGoogle Scholar | 29299268PubMed |

McComb, K., Shannon, G., Durant, S. M., Sayialel, K., Slotow, R., Poole, J., and Moss, C. (2011). Leadership in elephants: the adaptive value of age. Proceedings of the Royal Society B. Biological Sciences 278, 3270–3276.
Leadership in elephants: the adaptive value of age.Crossref | GoogleScholarGoogle Scholar |

Menon, V., and Tiwari, S. K. (2019). Population status of Asian elephants Elephas maximus and key threats. International Zoo Yearbook 53, 17–30.
Population status of Asian elephants Elephas maximus and key threats.Crossref | GoogleScholarGoogle Scholar |

Mikota, S. K. (2006). Integument system. In ‘Biology, medicine, and surgery of elephants’. (Eds M. E. Fowler and S. K. Mikota.) pp. 253–261. (Blackwell Publishing: Ames, IA, USA.)

Morfeld, K. A., Meehan, C. L., Hogan, J. N., and Brown, J. L. (2016). Assessment of body condition in African (Loxodonta africana) and Asian (Elephas maximus) elephants in North American zoos and management practices associated with high body condition scores. PLoS One 11, e0155146.
Assessment of body condition in African (Loxodonta africana) and Asian (Elephas maximus) elephants in North American zoos and management practices associated with high body condition scores.Crossref | GoogleScholarGoogle Scholar | 27416141PubMed |

Mumby, H. S., and Plotnik, J. M. (2018). Taking the elephants’ perspective: remembering elephant behavior, cognition and ecology in human–elephant conflict mitigation. Frontiers in Ecology and Evolution 6, 122.
Taking the elephants’ perspective: remembering elephant behavior, cognition and ecology in human–elephant conflict mitigation.Crossref | GoogleScholarGoogle Scholar |

Ngama, S., Korte, L., Bindelle, J., Vermeulen, C., and Poulsen, J. R. (2016). How bees deter elephants: Beehive trials with forest elephants (Loxodonta africana cyclotis) in Gabon. PLoS One 11, e0155690.
How bees deter elephants: Beehive trials with forest elephants (Loxodonta africana cyclotis) in Gabon.Crossref | GoogleScholarGoogle Scholar | 27196059PubMed |

Osborn, F. V. (2002). Oleoresin as an elephant repellent: field trials in the communal lands of Zimbabwe. The Journal of Wildlife Management 66, 674–677.
Oleoresin as an elephant repellent: field trials in the communal lands of Zimbabwe.Crossref | GoogleScholarGoogle Scholar |

Osborn, F. V. (2004). Seasonal variation in feeding patterns and food selection by crop-raiding elephants in Zimbabwe. African Journal of Ecology 42, 322–327.
Seasonal variation in feeding patterns and food selection by crop-raiding elephants in Zimbabwe.Crossref | GoogleScholarGoogle Scholar |

Pinter-Wollman, N. (2012). Human-elephant conflict in Africa: the legal and political viability of translocations, wildlife corridors, and transfrontier parks for large mammal conservation. Journal of International Wildlife Law and Policy 15, 152–166.
Human-elephant conflict in Africa: the legal and political viability of translocations, wildlife corridors, and transfrontier parks for large mammal conservation.Crossref | GoogleScholarGoogle Scholar |

Plotnik, J. M., Lair, R., Suphachoksahakun, W., and Waal, F. B. M. d. (2011). Elephants know when they need a helping trunk in a cooperative task. Proceedings of the National Academy of Sciences of the United States of America 108, 5116–5121.
Elephants know when they need a helping trunk in a cooperative task.Crossref | GoogleScholarGoogle Scholar | 21383191PubMed |

Pokharel, S. S., Seshagiri, P. B., and Sukumar, R. (2017). Assessment of season-dependent body condition scores in relation to faecal glucocorticoid metabolites in free-ranging Asian elephants. Conservation Physiology 5, cox039.
Assessment of season-dependent body condition scores in relation to faecal glucocorticoid metabolites in free-ranging Asian elephants.Crossref | GoogleScholarGoogle Scholar | 28721215PubMed |

Pokharel, S. S., Singh, B., Seshagiri, P. B., and Sukumar, R. (2019). Lower levels of glucocorticoids in crop-raiders: diet quality as a potential ‘pacifier’ against stress in free-ranging Asian elephants in a human-production habitat. Animal Conservation 22, 177–188.
Lower levels of glucocorticoids in crop-raiders: diet quality as a potential ‘pacifier’ against stress in free-ranging Asian elephants in a human-production habitat.Crossref | GoogleScholarGoogle Scholar |

Poole, J. H. (1989). Mate guarding, reproductive success and female choice in African elephants. Animal Behaviour 37, 842–849.
Mate guarding, reproductive success and female choice in African elephants.Crossref | GoogleScholarGoogle Scholar |

Poole, J. H., and Moss, C. J. (1989). Elephant mate searching: group dynamics and vocal and olfactory communication. Symposium of the Zoological Society of London 61, 111–125.

Prakash, T. G. S. L., Wijeratne, A. W., and Fernando, P. (2020). Human–elephant conflict in Sri Lanka: patterns and extent. Gajah 51, 16–25.

R Core Team (2020). ‘R: a language and environment for statistical computing.’ (R Foundation for Statistical Computing: Vienna, Austria.)

Rasmussen, H. B., Okello, J. B. A., Wittemyer, G., Siegismund, H. R., Arctander, P., Vollrath, F., and Douglas-Hamilton, I. (2008). Age- and tactic-related paternity success in male African elephants. Behavioral Ecology 19, 9–15.
Age- and tactic-related paternity success in male African elephants.Crossref | GoogleScholarGoogle Scholar |

Riddle, H. S., Schulte, B. A., Desai, A. A., and Meer, L. d. (2010). Elephants: a conservation overview. Journal of Threatened Taxa 2, 653–661.
Elephants: a conservation overview.Crossref | GoogleScholarGoogle Scholar |

Sach, F., Dierenfeld, E. S., Langley-Evans, S. C., Watts, M. J., and Yon, L. (2019). African savanna elephants (Loxodonta africana) as an example of a herbivore making movement choices based on nutritional needs. PeerJ 7, e6260.
African savanna elephants (Loxodonta africana) as an example of a herbivore making movement choices based on nutritional needs.Crossref | GoogleScholarGoogle Scholar | 30723615PubMed |

Sach, F., Yon, L., Henley, M. D., Bedetti, A., Buss, P., Boer, W. F. d., Dierenfeld, E. S., Gardner, A., Langley-Evans, S. C., Hamilton, E., Lark, R. M., Prins, H. H. T., Swemmer, A. M., and Watts, M. J. (2020). Spatial geochemistry influences the home range of elephants. The Science of the Total Environment 729, 139066.
Spatial geochemistry influences the home range of elephants.Crossref | GoogleScholarGoogle Scholar | 32388134PubMed |

Saif, O., Kansky, R., Palash, A., Kidd, M., and Knight, A. T. (2020). Costs of coexistence: understanding the drivers of tolerance towards Asian elephants Elephas maximus in rural Bangladesh. Oryx 54, 603–611.
Costs of coexistence: understanding the drivers of tolerance towards Asian elephants Elephas maximus in rural Bangladesh.Crossref | GoogleScholarGoogle Scholar |

Santiapillai, C., and Wijeyamohan, S. (2013). The first national survey of elephants in Sri Lanka. Current Science 105, 153–154.

Santiapillai, C., and Wijeyamohan, S. (2016). Conservation and the history of human−elephant relations in Sri Lanka. In ‘Conflict, negotiation, and coexistence: rethinking human−elephant relations in South Asia’. (Eds P. Locke and J. Buckingham.) pp. 229–241. (Oxford University Press: Oxford.)

Santiapillai, C., Silva, A., Karyawasam, C., Esufali, S., Jayaniththi, S., Basnayake, M., Unantenne, V., and Wijeyamohan, S. (1999). Trade in Asian elephant ivory in Sri Lanka. Oryx 33, 176–180.
Trade in Asian elephant ivory in Sri Lanka.Crossref | GoogleScholarGoogle Scholar |

Santiapillai, C., Wijeyamohan, S., Bandara, G., Athurupana, R., Dissanayake, N., and Read, B. (2010). An assessment of the human–elephant conflict in Sri Lanka. Ceylon Journal of Science. Biological Sciences 39, 21–33.
An assessment of the human–elephant conflict in Sri Lanka.Crossref | GoogleScholarGoogle Scholar |

Seltmann, M. W., Helle, S., Htut, W., and Lahdenperä, M. (2019). Males have more aggressive and less sociable personalities than females in semi-captive Asian elephants. Scientific Reports 9, 2668.
Males have more aggressive and less sociable personalities than females in semi-captive Asian elephants.Crossref | GoogleScholarGoogle Scholar | 30804421PubMed |

Shaffer, L. J., Khadka, K. K., Hoek, J. V. D., and Naithani, K. J. (2019). Human-elephant conflict: a review of current management strategies and future directions. Frontiers in Ecology and Evolution 6, 235.
Human-elephant conflict: a review of current management strategies and future directions.Crossref | GoogleScholarGoogle Scholar |

Shoshani, J., Adler, R., Baccala, M. J., Barry, S., Battiata, R., Bedore, M. P., Berbenchuk, S. A., Bielaczyc, R., Booth, G., Bozarth, N., Bulgarelli, M. A., Church, I., Cosgriff, J. W., Crowe, H., DeFauw, S. L., Denes, L., Efthyvoulidis, E., Ekstrom, M., Engelhard, J. G., English, P., Fairchild, D., Fischer, C., Frahm, K., Frederick, D., Fried, J., Gaskins, T., Gatt, J., Gentles, W., Goshgarian, H. G., Grabowski, S., Haase, D., Hajj, K., Hall, G., Hawkins, D., Heberer, C., Helinski, A., Henry, S. R., Heyka, C., Hurt, M., Kemppainen, M., Kendra, C., Koenig, J., Konarske, P., Konwinski, S., Kopacz, S., Lakits, V. T., Lash, S., Laughlin, D., Meyers, S., Mizeres, N. J., Morehead, K. M., Muraski, A., Murphy, S., Niedbala, J., Overbeck, G., Powitz, R., Rafols, J. A., Raymer, S. L., Rezzonica, L., Rossmoore, H. W., Sabo, D., Schwikert, P. J., Shy, E., Skoney, J., Smith, D., Spodarek, K. L., Sujdak, P. J., Tarrant, T., Thielman, R., Thielman, R., Tisch, F., Walowicz, L., Williams, J., and Yehiel, D. (1982). On the dissection of a female Asian elephant (Elephas maximus maximus Linnaeus, 1758) and data from other elephants. Elephant 2, 3–93.
On the dissection of a female Asian elephant (Elephas maximus maximus Linnaeus, 1758) and data from other elephants.Crossref | GoogleScholarGoogle Scholar |

Sitati, N. W., Walpole, M. J., Smith, R. J., and Leader-Williams, N. (2003). Predicting spatial aspects of human–elephant conflict. Journal of Applied Ecology 40, 667–677.
Predicting spatial aspects of human–elephant conflict.Crossref | GoogleScholarGoogle Scholar |

Smith, F. (1890). Histology of the skin of the elephant. Journal of Anatomy and Physiology 24, 493–503.
| 17231874PubMed |

Spearman, R. I. C. (1970). The epidermis and its keratinisation in the African elephant (Loxodonta africana). Zoologica Africana 5, 327–338.
The epidermis and its keratinisation in the African elephant (Loxodonta africana).Crossref | GoogleScholarGoogle Scholar |

Srinivasaiah, N. M., Anand, V. D., Vaidyanathan, S., and Sinha, A. (2012). Usual populations, unusual individuals: insights into the behavior and management of Asian elephants in fragmented landscapes. PLoS One 7, e42571.
Usual populations, unusual individuals: insights into the behavior and management of Asian elephants in fragmented landscapes.Crossref | GoogleScholarGoogle Scholar | 22916135PubMed |

Srinivasaiah, N., Kumar, V., Vaidyanathan, S., and Sukumar, R. (2019). All-male groups in Asian elephants: a novel, adaptive social strategy in increasingly anthropogenic landscapes of southern India. Scientific Reports 9, 8678.
All-male groups in Asian elephants: a novel, adaptive social strategy in increasingly anthropogenic landscapes of southern India.Crossref | GoogleScholarGoogle Scholar | 31273235PubMed |

Sukumar, R. (1989) ‘The Asian elephant: ecology and management.’ (Cambridge University Press: Cambridge.)

Sukumar, R. (1990). Ecology of the Asian elephant in southern India. II. Feeding habits and crop raiding patterns. Journal of Tropical Ecology 6, 33–53.
Ecology of the Asian elephant in southern India. II. Feeding habits and crop raiding patterns.Crossref | GoogleScholarGoogle Scholar |

Sukumar, R. (2003) ‘The living elephants: evolutionary ecology, behavior, and conservation.’ (Oxford University Press: Oxford.)

Sukumar, R., and Gadgil, M. (1988). Male–female differences in foraging on crops by Asian elephants. Animal Behaviour 36, 1233–1235.
Male–female differences in foraging on crops by Asian elephants.Crossref | GoogleScholarGoogle Scholar |

Sutradhar, B. C., Das, B. C., Ferdous, S., Rahman, M., Hossain, S., Hossain, F., and Hasanuzzaman, (2018). Penetrating gunshot wound, emphysema and inanition followedby death in a wild Asian elephant: a case report. International Journal of Avian & Wildlife Biology 3, 119–122.
Penetrating gunshot wound, emphysema and inanition followedby death in a wild Asian elephant: a case report.Crossref | GoogleScholarGoogle Scholar |

Tscharntke, T., Clough, Y., Wanger, T. C., Jackson, L., Motzke, I., Perfecto, I., Vandermeer, J., and Whitbread, A. (2012). Global food security, biodiversity conservation and the future of agricultural intensification. Biological Conservation 151, 53–59.
Global food security, biodiversity conservation and the future of agricultural intensification.Crossref | GoogleScholarGoogle Scholar |

van Aarde, R., Whyte, I., and Pimm, S. (1999). Culling and the dynamics of the Kruger National Park African elephant population. Animal Conservation 2, 287–294.
Culling and the dynamics of the Kruger National Park African elephant population.Crossref | GoogleScholarGoogle Scholar |

Varma, S., Baskaran, N., and Sukumar, R. (2012). Field key for elephant population estimation and age and sex classification: resource material for synchronized elephant population count using block count, line transect dung count method and waterhole count. Centre for Ecological Sciences, Indian Institute of Science, Bangalore, Karnataka, India.

Von Hagen, R. L., Kasaine, S., Githiru, M., Amakobe, B., Mutwiwa, U. N., and Schulte, B. A. (2021). Metal strip fences for preventing African elephant (Loxodonta africana) crop foraging in the Kasigau Wildlife Corridor, Kenya. African Journal of Ecology 59, 293–298.
Metal strip fences for preventing African elephant (Loxodonta africana) crop foraging in the Kasigau Wildlife Corridor, Kenya.Crossref | GoogleScholarGoogle Scholar |

Webber, C. E., Sereivathana, T., Maltby, M. P., and Lee, P. C. (2011). Elephant crop-raiding and human–elephant conflict in Cambodia: crop selection and seasonal timings of raids. Oryx 45, 243–251.
Elephant crop-raiding and human–elephant conflict in Cambodia: crop selection and seasonal timings of raids.Crossref | GoogleScholarGoogle Scholar |

Wiafe, E. D., and Sam, M. K. (2014). Evaluation of a low-tech method, pepper–grease, for combatting elephant crop-raiding activities in Kakum Conservation Area, Ghana. Pachyderm 55, 38–42.

Wickham, H., Averick, M., Bryan, J., Chang, W., McGowan, L. D. A., François, R., Grolemund, G., Hayes, A., Henry, L., Hester, J., Kuhn, M., Pedersen, T. L., Miller, E., Bache, S. M., Müller, K., Ooms, J., Robinson, D., Seidel, D. P., Spinu, V., Takahashi, K., Vaughan, D., Wilke, C., Woo, K., and Yutani, H. (2019). Welcome to the tidyverse. Journal of Open Source Software 4, 1686.
Welcome to the tidyverse.Crossref | GoogleScholarGoogle Scholar |

Williams, A. C., Johnsingh, A. J. T., and Krausman, P. R. (2001). Elephant–human conflicts in Rajaji National Park, northwestern India. Wildlife Society Bulletin 29, 1097–1104.

Wilson, S., Davies, T. E., Hazarika, N., and Zimmermann, A. (2015). Understanding spatial and temporal patterns of human–elephant conflict in Assam, India. Oryx 49, 140–149.
Understanding spatial and temporal patterns of human–elephant conflict in Assam, India.Crossref | GoogleScholarGoogle Scholar |

Wittemyer, G. (2011). Effects of economic downturns on mortality of wild African elephants. Conservation Biology 25, 1002–1009.
Effects of economic downturns on mortality of wild African elephants.Crossref | GoogleScholarGoogle Scholar | 21790785PubMed |

Woodroffe, R., Thirgood, S., and Rabinowitz, A. (2005) ‘People and wildlife: conflict or coexistence?’ (Cambridge University Press: Cambridge.)

Yasui, S., Konno, A., Tanaka, M., Idani, G. i., Ludwig, A., Lieckfeldt, D., and Inoue-Murayama, M. (2013). Personality assessment and its association with genetic factors in captive Asian and African elephants. Zoo Biology 32, 70–78.
Personality assessment and its association with genetic factors in captive Asian and African elephants.Crossref | GoogleScholarGoogle Scholar | 22996044PubMed |

Zimmermann, A., Davies, T. E., Hazarika, N., Wilson, S., Chakrabarty, J., Hazarika, B., and Das, D. (2009). Community-based human-elephant conflict management in Assam. Gajah 30, 34–40.