As charismatic megafauna, flagship species and ecosystem engineers, African elephants are vital to African tourist economy and ecosystem health. Wildlife tourism may induce behavioural shifts that push desired animals into less frequented areas, disrupt natural behaviours and potentially have negative ecosystem consequences. Managers of tourism areas need to make decisions about the level and distribution of tourism within their managed area, such as a national park, to determine the best combination of providing a memorable tourist experience and ensuring the wellbeing of the species sought by tourists, such as elephants and their habitat. To assess this trade-off, we examined whether African elephants are less vigilant or more vigilant in areas with lower levels of human traffic. We studied elephant behaviours in Zambezi National Park near Victoria Falls, Zimbabwe. Geographical, demographic and behavioural data were collected via elephant sightings and any vehicles in the area. We mapped these data in ArcGIS to create a visual representation of their spatial relationship and to identify hotspot locations. Elephants were more frequently sighted in the park region with less vehicle traffic, but they also displayed higher rates of vigilance behaviours in that region, suggesting greater vigilance where vehicles were less common. An understanding of elephant behaviour even within managed areas can inform decisions on tourism levels across locations to provide elephants with areas of refuge from human intrusion, but additional research is needed to determine the longer-term ecosystem consequences. (c) 2026 The Author(s). Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/ licenses/by-nc-nd/4.0/).

ABSTRACT Elephant behaviour towards dead conspecifics has been well documented, but mostly on fresh female or calf carcasses. This study builds upon existing observations by describing elephant interactions at the mortality site of a long‐dead bull African savannah elephant in southern Kenya. A camera was installed at the site for 3.5 months in 2023, during which 34 unique visits were recorded. Visiting elephants often interacted with (e.g., touched, sniffed) the bones. Most events (85.3%) were by lone males or bachelor groups. Patterns of visitation are compared to past studies, and hypotheses explaining the prolonged visitation to dead conspecifics are discussed.

Ecotourism promises to reconcile wildlife conservation and human development if negative impacts of human visitation and associated infrastructure can be minimized. Animal behavior studies can be used to identify individual and population responses to anthropogenic impacts before other fitness consequences are documented. With input from professionals in animal behavior and ecotourism, we identified key questions needed to better understand the impact of ecotourism on wildlife. Activity budgets, foraging, movement, stress, habituation, and reproduction were themes that emerged from our survey. We highlight promising research on these themes and identify remaining behavioral research questions about conserving wildlife in the context of ecotourism. Although ecotourism activities often have detrimental effects on animal behavior, we highlight research needs that can inform management and ecotourist education to improve human behavior to be more compatible with sustainable use of nature.

Elephant deterrent fences have become a popular strategy to mitigate human-elephant conflict in agricultural areas. It is unknown whether the adoption of climate-smart agricultural practices like intercropping could influence elephant behaviour and therefore the effectiveness of these fences. In this study in southern Kenya, the success of the Kasaine metal strip fence was compared between intercropped and monocropped fields. Results support previous research, as the Kasaine fence remained effective, suggesting the adoption of intercropping is unlikely to drastically change elephant crop foraging behaviour.

Anthropogenic stressors, such as habitat loss and fragmentation, destabilize animal populations, putting species at risk of extirpation or extinction. Conservation translocations, i.e. moving animals from one area of their native habitat to create new populations or bolster existing populations in other areas, are increasingly necessary to combat species decline. However, translocation success is variable, and failure is common, regardless of the resources invested in the programme. Additional anthropogenic stressors, such as climate change, pollution, harvesting, invasive species and human-wildlife conflict, further complicate the translocation process. Methods to reduce variable results and increase success are needed, and incorporating animal behaviour and its relevant theories has that potential. We reviewed cases where animal behaviour contributed to the success or failure of a translocation programme where anthropogenic stressors (in particular, climate change, pollution, harvesting, invasive species and human-wildlife conflict) contributed to or were the driving force of the programme's outcome. We further identify areas where conservation behaviour could be implemented to improve the success of translocation programmes and highlight why we need to better leverage our knowledge of animal behaviour to improve the success of translocations in a rapidly changing environment. (c) 2025 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights are reserved, including those for text and data mining, AI training, and similar technologies.

African elephants (Loxodonta africana) play a significant role in the modification of their habitat, foremost by decreasing woody vegetation cover and density. Little is known regarding the effects of elephant habitat disturbance (EHD) on medium-to-large mammal and bird communities. While the intermediate disturbance hypothesis (IDH) predicts species richness and diversity will be maximised in areas of moderate EHD, high levels of EHD could result in habitat degradation and negatively impact certain wildlife communities. To examine this relationship, we compared mammal and avian community assemblages, richness, and diversity across differing severities (low, medium, high) of elephant disturbed habitats (woodland, shrubland, and mixed wood/shrubland) within Rukinga Wildlife Sanctuary (RWS) in south-eastern Kenya. From June-November 2022, we collected wildlife detection data via driving transects and surveyed EHD through observation of elephant tree damage. While EHD level did impact community assemblage, possibly through vegetation modification and/or increased habitat heterogeneity, EHD had no negative impacts on species richness and diversity in the three habitat types. In contrast to our IDH prediction, richness and diversity were highest in high EHD areas within shrubland habitat and showed no difference across EHD levels for the other two habitats. Overall, this study provides evidence that elephants in RWS are not disturbing habitats to an extent that negatively impacts sanctuary viability, and at least for shrub habitat appear to enhance it. Additionally, smaller wildlife habitats may be able to mitigate lasting elephant overpopulation damages through increased connectivity to other protected areas.

Crop foraging by African savannah elephants Loxodonta africana negatively affects farmer livelihoods and support for conservation, yet affordable, sustainable and practical solutions remain elusive. To inform conservation priorities, our goal was to assess the hitherto little explored relationships between farmers’ views on agricultural damage and the socio-economic factors limiting their use of elephant deterrents. We tested our hypotheses associated with the demographic categories of age, education level, years spent farming, gender, exposure to information on deterrent methods, farm size, village and relevant combinations of these factors by surveying 206 respondents across six villages in rural Kenya and analysing the resulting data using an information theoretic approach. Respondents were almost four times more likely to use deterrents if exposed to the relevant information, and almost five times more likely to do so if they had secondary education as opposed to none. Farmers with a higher level of education were five times more likely to have received information on deterrents compared to those with no formal education. Participants who had not received information on deterrents were almost three times more likely to believe that they could implement deterrent methods. Respondents who stated that they could not implement deterrents overwhelmingly cited a lack of financial resources as the reason. Overall, we found that crucial information on reducing elephant crop foraging is not reaching the relevant stakeholders, and socio-economic factors such as education and exposure to information appear to limit uptake of protective measures. These insights are important for developing mitigation strategies and supporting the livelihoods of people affected by negative human–elephant interactions, and thus for effective elephant conservation. Our findings also have broader applications for practitioners seeking to understand barriers stakeholders face in their efforts to mitigate negative interactions with wildlife.

Horses can become obese and develop related health issues such as laminitis from excessive grazing on high-quality pasture grass; limiting pasture intake can allow weight loss to occur. The objective of this study was to determine the effect of space-restricted rotational grazing on body weight (BW) and time budgets in horses. Eight mature geldings and mares with maintenance-only requirements were randomly assigned to either a space-restricted rotational grazing group (SRG; BW 512 ± 6 kg; n = 4) or a continuous grazing group (CG; BW 517 ± 49 kg; n = 4) for 42 d SRG horses grazed an area with dimensions to provide 80–90 % of mean digestible energy requirement for the 4 horses over a 7-d grazing period; whereas, the CG horses continuously grazed similar non-toxic endophyte-infected tall fescue pasture providing greater than maintenance requirements for the 42 d Horses in the SRG group were moved to a new area every 7 d for 6 weeks. On d 7 at 1600 h of each week, horses were brought inside, and feed was withheld overnight. At 0700 h the next day, BWs were recorded prior to turnout. Observers recorded behaviors simultaneously on SRG and CG horses every six minutes throughout the day three days per week according to an ethogram. This included 30 s scans of all horses. Proportion of grazing and standing had an inverse relationship. Proportion of grazing was affected by the treatment by time interaction, which grazing was displayed more in SRG than CG during weeks 2 and 3, and then reversed weeks 4, 5 and 6.

Risk assessment relies on cost-benefit analysis by prey species in relation to predator behavior. The lunar cycle can have strong influences on the way that predator and prey species behave in their environment as light levels can influence activity patterns. By examining how lunar phase influences behavior, we may be able to better predict instances of human-wildlife conflict. As part of the Elephants and Sustainable Agriculture in Kenya project, over 5 years of study have been dedicated to mitigating human-elephant conflict in the Kasigau Wildlife Corridor in southern Kenya, focusing on the construction of deterrent fences and the collection of environmental data in and bordering the Rukinga Wildlife Sanctuary. Elephant presence around crop fields was hypothesized to have a negative relationship with lunar light levels because of an increase in perceived risk from typically greater human activity with moonlight, and a positive relationship with increased darkness during the waning moon phases. The lunar phase data were compared to elephant presence within 12 m of experimental crop fields along the boundary of their wild habitat and human agricultural land to provide insight for potential crop raiding events. Elephants were present significantly less during the full and gibbous moon phases when compared to the new moon phase, and significantly more during the new and waning moon phases. Our findings are being shared with the farmers living in the KWC and may be useful to others living in high HEC areas by providing a form of early warning system that also may have broader applications in the mitigation of other human-wildlife interactions. HIGHLIGHTSDynamic landscape of fear provides novel insight on elephant behavior near agricultural lands with applications for conservation strategies.

Negative human-wildlife interactions are a growing problem, particularly for people living near protected areas and wildlife refuges. In Kenya, African savannah elephants Loxodonta africana threaten food security for subsistence farmers by crop foraging, which can jeopardize conservation efforts if farmers retaliate against elephants. To inform conservation and management, this study had three objectives: (1) to evaluate stakeholder participatory models of human-elephant conflict; (2) to note any novel or under-represented variables in the models; and (3) to deter-mine if there were indicators for assessing the success of mitigation programmes using a biocultural approach. We conducted participatory modelling sessions in six villages in rural Kenya using fuzzy cognitive mapping (n =206 participants). Farmers created group visual models with variables related to conflict with elephants. A total of 14 variables were common across all six villages, with the two highest centrality scores (a measure of importance to overall dynamics) associated with income and feelings of security. Most variables fell into two categories: environmental interactions, and policy and management. Multiple variables such as road infrastructure (drivers) and soil compaction (con-sequences) were identified as aspects of conflicts that are under-reported or absent in scientific literature, as well as potential socio-cultural indicators. The participatory method used is a tool for gaining more refined insights into inter-actions with elephants, with implications for other complex conservation issues or wildlife interactions. A more holistic view of the impacts of human-elephant interactions as demonstrated here can lead to sustainable, co-developed pro-grammes that benefit both farmer livelihoods and elephant conservation.

In semi-arid southern Kenya, climate change is putting rainfed agriculture at risk with major implications for food security. The zai pit, a Climate-Smart Agriculture (CSA) technique, has proven to enhance yields in arid regions, but its labor demands have limited adoption rates. This study assessed how the zai pit depth and manure application within zai pits influenced maize (Zea mays) success (i.e., growth, development, and productivity). Three zai pit treatments were prepared at the Wildlife Works Research Center in southeastern Kenya: (1) deep (50 cm) with manure; (2) shallow (25 cm) with manure; and (3) deep (50 cm) without manure, and all were compared to a non-zai pit control (surface planting). Maize growth/development (e.g., height, stage, roots) and productivity (e.g., yield) measurements were taken over two growing periods. For most measures, shallow zai pits performed equally as well as deep zai pits, with both performing better than the control. Zai pits without manure performed significantly worse than zai pits with manure, oftentimes not differing from the control. Results suggest that maize success is influenced more by manure than the depth of the pit. Kenyan farmers are encouraged to dig shallower, manure-enriched zai pits to enhance food security in response to climate change.

Interactions with elephants are increasing throughout the African savannah elephant range as habitat loss and modification and a growing human population continue to bring people and elephants into contact. These interactions can become negative when elephants are tempted into farmland and consume farmers’ crops or destroy water supplies, leading to human-elephant conflict. Each community coping with regular wildlife conflicts faces unique socio-ecological circumstances and constraints. Therefore, understanding the challenges within each system is a crucial step in designing customised management plans and mitigation interventions, adapted to the situation. We used a combination of survey results and participatory group fuzzy logic cognitive maps from six farming communities in south-east Kenya from previous studies, to understand the complex drivers and consequences of conflicts with elephants and how farmers conceptualize these interactions. These data informed the creation of five main strategies for mitigating the impacts of crop-raiding by elephants: deterrent methods, climate-smart agricultural techniques, alternative livelihoods, safety around elephants, and environmental stewardship. We consulted with local experts to design and deliver workshops in the six communities to present potential solutions within the strategies and to provide the content for a take-home manual. Although no single solution has emerged as the ideal way to mitigate these encounters, the workshops demonstrated a variety of approaches that can alleviate the financial and safety concerns of farmers. Future work should include understanding barriers to wider acceptance of such methods and evaluating the efficacy of multifaceted approaches. Creating a customised curriculum for workshops informed by social science data can provide vital information for local people who want to co-exist alongside elephants and other wildlife. Additional keywords: community-based conservation, conservation planning, conservation social science, human-wildlife co-existence

Human-elephant conflict (HEC) compromises crop security and threatens elephant conservation. Most commonly, HEC manifests as crop-foraging as elephants modify natural foraging strategies to incorporate crops. Farmers may retaliate by frightening or harming elephants, leaving scars from inflicted wounds. We assessed the prevalence and distribution of scars on the bodies of African savanna elephants (Loxodonta africana) observed in the Kasigau Wildlife Corridor (KWC), part of the Greater Tsavo Ecosystem of Kenya, where conflict is prevalent. We surmised that scars on the body are largely a result of HEC as opposed to scars on the rump or head, which we attributed primarily to elephant-elephant conflict. We hypothesized that: (1) male elephants would have more scars than females; (2) older males would be more likely to have scars than younger males; and (3) most scars would be located on the bodies of elephants. We assessed scars from a photographic catalogue of elephants from the KWC. In line with our hypotheses, male elephants were more likely to have scars than females (32% of males compared to 6% of females); older males had significantly more scars than younger males (61% compared to 24%); and the majority of scars (89%) were located on the body. Scar presence may be useful as an animal-centered indicator to estimate the prevalence and demographic patterns of HEC.

As human–elephant coexistence is challenged across Africa and Asia, conservationists seek methods to mitigate elephant crop raiding. Farmers rely on their crops for sustenance, and crop raiding events can destroy harvests. To mitigate these conflicts, deterrent fences continue to be developed. Modern deterrent techniques incorporate improved understanding of elephant biology, including their reliance on chemical signals. The construction of deterrent fences with cloths soaked in chili pepper extracts with high Scoville heat units (SHU) and used engine oil has become widespread but with variable effectiveness. However, the active levels of capsaicinoids in these mixtures and the rate of degradation of these compounds remain largely unexplored. In Kenya, our field experiments found that chili fences performed less effectively than desired possibly due to environmental conditions. In the lab, we evaluated SHU from chili peppers in three preparation recipes (vodka, water, and oil) and found that mixing with oil was crucial for increased solubility. We simulated a chili fence in the lab and detected no significant SHU loss over 7 days, which has positive implications for fence maintenance. Continuous exploration of the applications of chemical ecology to major conservation issues such as crop raiding can benefit humans and wildlife globally.

Farmers in sub-Saharan Africa are facing threats to their livelihoods including climate change, unsustainable farming practices, increasing water scarcity, and crop raiding herbivores, among other challenges. How demographic, economic, and social factors impact farmers’ attitudes and responses to such threats and the relationship to conservation initiatives remains an underexplored topic. With the aim of informing policy and management priorities locally, but with broad applications, we developed a set of a priori hypotheses related to the impacts of livelihood threats and alternatives for farmers. To assess these hypotheses, we surveyed 206 farmers across 6 villages in the Kasigau Wildlife Corridor of Kenya. Most respondents believed that climate change had negatively affected their lives and a third of respondents had not been exposed to alternative crop types. However, sociodemographic factors were poor predictors of farmer beliefs and knowledge, suggesting a homogeneity of views. Nearly half (48%) of respondents preferred to continue farming despite other possible means of income, potentially indicating a strong sense of place. Survey questions revealed other disparities such as 55% of respondents had no formal farm training outside of school, and most had never visited the nearby national parks. Compared to other studies, an above average belief in the benefits of wildlife was notable across the villages. Our findings can inform programs that work with local farmers to create specific management priorities that assist stakeholders in building resilience to threatened natural resources and increase food security which could bolster support of conservation programs.

Group-living animals that live in complex social systems require effective modes of communication to maintain social cohesion, and several acoustic, olfactory and visual signaling systems have been described. Individuals need to discriminate between in- and out-group odour to both avoid inbreeding and to identify recipients for reciprocal behaviour. The presence of a unique group odour, identified in several social mammals, is a proposed mechanism whereby conspecifics can distinguish group from non-group members. African elephants ( Loxodonta africana ) live in stable, socially complex, multi-female, fission–fusion groups, characterized by female philopatry, male dispersal and linear dominance hierarchies. Elephant social behaviour suggests that individuals use odour to monitor the sex, reproductive status, location, health, identity and social status of conspecifics. To date, it is not clear what fixed or variable information is contained in African elephant secretions, and whether odour encodes kinship or group membership information. Here we use SPME GC–MS generated semiochemical profiles for temporal, buccal and genital secretions for 113 wild African elephants and test their relationship with measures of genetic relatedness. Our results reveal the existence of individual identity odour profiles in African elephants as well as a signature for age encoded in temporal gland and buccal secretions. Olfactory signatures for genetic relatedness were found in labial secretions of adult sisters. While group odour was not correlated with group genetic relatedness, our analysis identified “group membership” as a significant factor explaining chemical differences between social groups. Saturated and short-chain fatty acids (SCFAs), derived from key volatile compounds from bacterial fermentation, were identified in temporal, buccal and genital secretions suggesting that group odour in African elephants may be the result of bacterial elements of the gut microbiome. The frequent affiliative behavior of African elephants is posited as a likely mechanism for bacterial transmission. Our findings favour flexible group-specific bacterial odours, which have already been proposed for other social mammals and present a useful form of olfactory communication that promotes bond group cohesion among non-relatives in fission–fusion mammals.

Sexual selection mediated by multimodal signals is common among polygynous species, including seasonally breeding mammals. Indirect benefit models provide plausible explanations for how and why mate selection can occur in the absence of direct benefits. Musth - an asynchronous reproductive state in male elephants - facilitates both inter- and intrasexual selection via indirect benefits, and it is further communicated through a multimodal signal. In this review, we synthesise existing evidence that supports the hypothesis that musth is a multimodal signal subject to sexual selection and that male elephants increase their direct fitness by propagating this signal while females accrue indirect benefits. Musth is characterised by a suite of physiological and behavioural changes, serving to facilitate copulation between the sexes, and via multisensory modalities musth conveys honest information about the condition of a male. Female elephants mate preferentially with musth males, increasing their own fitness in the absence of direct benefits. In addition, musth for costly physical combat. Future work in this field should investigate potential postcopulatory selection mechanisms in elephants, including sperm competition and cryptic female choice. These topics join other fundamental questions related to sexual selection, signalling, and indirect benefits that are still unanswered in elephants.

The use of odors is a popular form of sensory enrichment, yet few studies have investigated the longer lasting effects of biologically relevant chemical signals like pheromones. Asian elephants (Elephas maximus) are unique because they are commonly held in zoos and utilize two single-compounds to synchronize reproduction: frontalin (a pheromone produced by male elephants in musth) and (Z)-7-dodecenyl acetate (Z7-12:Ac, a pheromone produced by female elephants around ovulation). While Asian elephant chemical signaling dynamics have been well-studied, no reports exist on the broader behavioral effects (i.e., primer effects) of either of these pheromones. The purpose of this study was to describe the primer effects of both frontalin and Z7-12:Ac for potential enrichment applications. We bioassayed 50 elephants at 10 zoos using synthetic versions of these two pheromones that replicated the concentrations at which they are produced naturally. We found that behavioral diversity in response to these pheromones was influenced both by the degree of exposure to the chemicals and by properties of the receiver, such as sex, sexual experience, age, and social access, with models that included all or most of these factors explaining much of the variation in the bioassay data for frontalin (R-(c)(2) = 0.42) and Z7-12: Ac (R-(c)(2) = 0.59). Furthermore, both males and females exhibited distinct changes in activity levels in the presence of frontalin or Z7-12:Ac, with specific behavioral differences noted. For instance, frontalin stimulated increased activity (and decreased inactivity) among female elephants (P < 0.001). Foraging (P = 0.029) and walking (P = 0.001) increased when males were exposed to frontalin, but it also stimulated increased time spent engaged in stationary stereotypy (P < 0.001). On the other hand, Z7-12:Ac encouraged greater activity levels in female (P < 0.001) and male (P = 0.010) elephants. These results support the use of pheromones as enrichment opportunities for elephants, and suggest appropriate chemical signals should be considered for other species. Because these compounds hold biological, ecological, and/or evolutionary salience, they can have long-lasting effects and are particularly relevant to long-term management strategies. In conjunction with other forms of sensory enrichment-including biologically novel odors-pheromones have the potential to enhance the wellness of zoo-housed species.

Chemical signals are the oldest and most ubiquitous means of mediating intra- and interspecific interactions. The three extant species of elephants, the Asian elephant and the two African species, savanna and forest share sociobiological patterns in which chemical signals play a vital role. Elephants emit secretions and excretions and display behaviors that reveal the importance of odors in their interactions. In this review, we begin with a brief introduction of research in elephant chemical ecology leading up to the 21st century, and then we summarize the body of work that has built upon it and occurred in the last c. 20 years. The 21st century has expanded our understanding on elephant chemical ecology, revealing their use of odors to detect potential threats and make dietary choices. Furthermore, complementary in situ and ex situ studies have allowed the careful observations of captive elephants to be extended to fieldwork involving their wild counterparts. While important advances have been made in the 21st century, further work should investigate the roles of chemical signaling in elephants and how these signals interact with other sensory modalities. All three elephant species are threatened with extinction, and we suggest that chemical ecology can be applied for targeted conservation efforts.