Animals living in heterogeneous landscapes are often faced with making a trade-off between maximizing foraging success and avoiding risk. Using high-resolution GPS-tracking data, this study explored the fine-scale movement patterns and risk sensitivity of crop-raiding African elephants, Loxodonta africana, in the anthropogenic landscape of Tsavo, Kenya. We analysed patterns in the speed and tortuosity of elephant movements over the 24 h surrounding crop-raiding events and compared them with those of nonraiding elephants during corresponding periods. Crop-raiding elephants moved faster and straighter (less tortuously) with closer temporal proximity to farmland, which we argue reflects their increased intensity of risk avoidance behaviours in response to approaching humans. Once inside farmland, elephants appeared to reduce movements associated with risk avoidance to forage intensively on crops, decreasing their speed and reducing the likelihood of moving in straight lines while crop raiding. These results highlight trade-offs in the fine-scale movement patterns of elephants living in anthropogenic landscapes with differing levels of habitat quality and exposure to humans, providing new insight into how they perceive the risks associated with crop raiding.

Abstract As human‐elephant conflict (HEC) increases, a better understanding of the human dimensions of these conflicts and non‐violent mitigation methods are needed to foster long‐term coexistence. In this study, we conducted household questionnaires (n = 296) to assess the prevalence of HEC and attitudes towards elephants in four rural villages in Thailand. In addition, we evaluated a pilot beehive fence as a sustainable solution for HEC. The majority of the households reported seeing or hearing elephants near their property at least once a week (84.9%) and experienced negative impacts from elephants in the last 5 years, (81.0%). The beehive fence deterred 88.4% of individual elephants (n = 155) and 64.3% of elephant groups (n = 28) that approached the fence. Most elephants (70.7%) exhibited behaviors suggesting heightened attentiveness or alarm. The farm owner reported economic and social benefits of the beehive fence. By contributing to farmer income and reducing crop damage caused by wild elephants, beehive fencing may provide an important locally‐managed complement to regional HEC mitigation methods.

Human-elephant conflict (HEC) in the form of crop-raiding, is a major conservation challenge to the long-term survival of elephant populations, simultaneously threatening the livelihoods and personal safety of people living in proximity to elephants. The widespread problem of HEC has led to a great deal of research into the causes, consequences and predictors of elephant crop-raiding activity. However, despite similarities across HEC situations, site-specific differences are also apparent. Furthermore, most studies focus on one facet of HEC when it is a complex issue requiring understanding of local elephant behaviour, identification of the characteristics and patterns of cropraiding at the local scale, and careful implementation and monitoring of mitigation strategies. In this study, I selected a region of Sri Lanka experiencing high levels of HEC and sought to provide an in-depth assessment of the site-specific situation generated over a three-year period. Specifically, we aimed to: identify general patterns of behaviour occurring in local areas representing differing levels of anthropogenic disturbance to elephants; profile patterns and predictors of crop-raiding activity in a village heavily impacted by HEC; and test the effectiveness of beehive fencing as an Asian elephant deterrent tool. First, I provide initial evidence that elephants inhabiting areas of ‘medium’ level anthropogenic disturbance outside of protected boundaries, interrupt feeding and increase ‘reactive’ behaviours such as smelling and holding the ‘vigilance’ posture in response to immediate anthropogenic threats in the environment. In the absence of any known disturbances, there was no difference in general behaviours of male or female elephants between the two risk zones. Secondly, I show that elephant crop-raiding in Dewagiriya Village occurs year-round and follows no clear seasonal patterns. Similar to other HEC situations, male elephants are the predominant crop-raiders, and crop-raiding occurs almost exclusively at nights. Within-site variations in crop-raiding intensity were also identified, with properties closest to water tanks and forest habitat the most vulnerable. Finally, our three-year beehive fence trial showed that households using beehive fences around their gardens had significantly less elephant visits into their gardens then households without. Still, difficulties in attracting natural colonies, poor honey production, set-up costs, and farmer motivations were barriers to success. This study contributes to the general body of knowledge on elephant behaviour in anthropogenically influenced contexts, and specifically on patterns of crop-raiding and mitigation efforts. Further research into the potential of beehive fences as an Asian elephant deterrent, preferably in a location more amenable to beekeeping, would help to determine the value of expanding this technique further in Sri Lanka, and elsewhere in Asia.

Landscape planning that ensures the ecological integrity of ecosystems is critical in the face of rapid human‐driven habitat conversion and development pressure. Wildlife tracking data provide unique and valuable information on animal distribution and location‐specific behaviors that can serve to increase the efficacy of such planning. Given the spatiotemporal complexity inherent to animal movements, the interaction between movement behavior and a location is often oversimplified in commonly applied analyses of tracking data. We analyzed GPS‐tracking‐derived metrics of intensity of use, structural properties (based on network theory), and properties of the movement path (speed and directionality) with machine learning to define homogeneous spatial movement types. We applied our approach to a long‐term tracking data set of over 130 African elephants (Loxodonta africana ) in an area under pressure from infrastructure development. We identified 5 unique location‐specific movement categories displayed by elephants, generally defined as high, medium, and low use intensity, and 2 types of connectivity corridors associated with fast and slow movements. High‐use and slow‐movement corridors were associated with similar landscape characteristics associated with productive areas near water, whereas low‐use and fast corridors were characterized by areas of low productivity farther from water. By combining information on intensity of use, properties of movement paths, and structural aspects of movement across the landscape, our approach provides an explicit definition of the functional role of areas for movement across the landscape that we term the movescape . This combined, high‐resolution information regarding wildlife space use offers mechanistic information that can improve landscape planning.

Projects Updates from Save the Elephants

Abstract Wildlife counts in Africa and elsewhere are often implemented using light aircraft with ‘rear-seat-observer’ (RSO) counting crews. Previous research has indicated that RSOs often fail to detect animals, and that population estimates are therefore biased. We conducted aerial wildlife surveys in Murchison Falls Protected Area, Uganda, in which we replaced RSOs with high-definition ‘oblique camera count’ (OCC) systems. The survey area comprises forests, woodlands and grasslands. Four counts were conducted in 2015–2016 using a systematic-reconnaissance-flight (SRF) strip-transect design. Camera inclination angles, focal lengths, altitude and frame interval were calibrated to provide imaged strips of known sample size on the left and right sides of the aircraft. Using digital cameras, 24 000 high-definition images were acquired for each count, which were visually interpreted by four airphoto interpreters. We used the standard Jolly II SRF analysis to derive population estimates. Our OCC estimates of the antelopes – hartebeest, Uganda kob, waterbuck and oribi – were, respectively, 25%, 103%, 97% and 2100% higher than in the most recent RSO count conducted in 2014. The OCC surveys doubled the 2014 RSO estimate of 58 000 Uganda kob to over 118 000. Population size estimates of elephants and giraffes did not differ significantly. Although all four OCC buffalo estimates were higher than the RSO estimates – in one count by 60% – these differences were not significant due to the clumped distribution and high variation in herd sizes, resulting in imprecise estimation by sampling. We conclude that RSO wildlife counts in Murchison have been effective in enumerating elephants and giraffe, but that many smaller species have not been well detected. We emphasize the importance of 60 years of RSO-based surveys across Africa, but suggest that new imaging technologies are embraced to improve accuracy.

Abstract Conflict between humans and elephants is a notoriously complex problem requiring a detailed understanding of the underlying patterns and processes in order to develop effective solutions. Advances in radio tracking technologies have enabled researchers to examine in detail the ways in which tracked animals interact with their environments. We analysed the movement patterns of an habitual crop raiding African elephant (Loxodonta Africana) in the Amboseli ecosystem in southern Kenya. We identified three distinct patterns of movement associated with instances of crop raiding; these were (1) opportunistic raiding, (2) purposeful raiding, and (3) incidental raiding. The distinct characteristics of each of these movements serve to demonstrate the differing circumstances under which elephants are brought into contact with agricultural areas in their daily negotiations of the network of human land-use and protected areas. These findings highlight the need to understand patterns of elephant movement and interactions with farmland in order to craft management strategies that are effective in reducing levels of human-elephant conflict and promote tolerance of elephants in rural communities.

Abstract In aerial wildlife counts, human observers often fail to detect animals. We conducted a multi-species sample-count in Tsavo National Park, Kenya, with traditional rear-seat-observers (RSOs) and an automated ‘oblique-camera-count’ (OCC) imaging system to compare estimates of 23 wildlife species derived from these two survey methods. An aerial Total Count of elephant, buffalo and giraffe, conducted a month previously, provided a further comparison. In the Tsavo Core (9560 km2), which harbours 80% of Tsavo’s elephants, the OCC system acquired 81 000 images for interpretation, of which 67 000 were obtained in parallel with RSO-counting along 3004 km of flight line. The Tsavo outer blocks (24 171 km2) were surveyed using the OCC system without RSOs to acquire a further 84 000 images. A random sample of 11 553 images were re-interpreted to derive species-specific probabilities of detection and correction factors. Using ‘Jolly II’, non-parametric and Bayesian analyses, and applying correction factors, we demonstrate that the RSOs did not detect 14% of elephants, 60% of giraffe, 48% of zebra and 66% of the large antelopes. For comparison, the Total Count observers did not detect 27% of elephant, 33% of buffalo, 57% of giraffe and 85% of carcasses. The OCC method raises the elephant population estimate to 16 681 ± 4047 (95% cl) from the 12 722 counted in the Total Count (Z = 1.917, p = .0276). These results suggest that RSO-based methods have significantly undercounted wildlife populations. To align with improved counting methods, previous results need to be re-calibrated.

Abstract Natural habitats are rapidly being converted to cultivated croplands, and crop-raiding by wildlife threatens both wildlife conservation and human livelihoods worldwide.  We combined movement data from GPS-collared elephants with camera-trap data and local reporting systems in a before–after-control-impact design to evaluate community-based strategies for reducing crop raiding outside Mozambique’s Gorongosa National Park. All types of experimental fences tested (beehive, chili, beehive and chili combined, and procedural controls) significantly reduced the number of times elephants left the Park to raid crops. However, placing beehive fences at a subset of key crossing locations reduced the odds that elephants would leave the Park by up to 95% relative to unfenced crossings, and was the most effective strategy. Beehive fences also created opportunities for income generation via honey production. Our results provide experimental evidence that working with local communities to modify both animal behavior and human attitudes can mitigate conflict at the human–wildlife interface.

Abstract 1. Long‐term bio‐logging has the potential to reveal how movements, and hence lifehistory trade‐offs, vary over a lifetime. Reproductive tactics in particular may vary as individuals' trade‐off current investment versus lifetime fitness. Male African savanna elephants (Loxodona africana) provide a telling example of balancing body growth with reproductive fitness due to the combination of indeterminate growth and strongly delineated periods of sexual activity (musth), which results in reproductive tactics that alter with age. 2. Our study aims to quantify the extent to which male elephants alter their movement patterns, and hence energetic allocation, in relation to (a) reproductive state and (b) age, and (c) to determine whether musth periods can be detected directly from GPS tracking data. 3. We used a combination of GPS tracking data and visual observations of 25 male elephants ranging in age from 20 to 52 years to examine the influence of reproductive state and age on movement. We then used a three‐state hidden Markov model (HMM) to detect musth behaviour in a subset of sequential tracking data. 4. Our results demonstrate that male elephants increased their daily mean speed and range size with age and in musth. Furthermore, non‐musth speed decreased with age, presumably reflecting a shift towards energy acquisition during non‐musth. Thus, despite similar speeds and marginally larger ranges between reproductive states at age 20, by age 50, males were travelling 2.0 times faster in a 3.5 times larger area in musth relative to non‐musth. The distinctiveness of musth periods over age 35 meant the three‐state HMM could automatically detect musth movement with high sensitivity and specificity, but could not for the younger age class. 5. We show that male elephants increased their energetic allocation into reproduction with age as the probability of reproductive success increases. Given that older male elephants tend to be both the target of legal trophy hunting and illegal poaching, man‐made interference could drive fundamental changes in elephant reproductive tactics. Bio‐logging, as our study reveals, has the potential both to quantify mature elephant reproductive tactics remotely and to be used to institute proactive management strategies around the reproductive behaviour of this charismatic keystone species.

ABSTRACT Poaching is the most immediate threat to African elephants (Loxodonta africana). Several continental‐wide surges in poaching have occurred since the latter half of the twentieth century, and the latest surge occurred from 2007 to 2012. The behavioral responses of elephants to poaching risk has not been studied widely because of a lack of high‐resolution movement data collected simultaneously with verified causes of mortality. We managed to collate 2 such datasets from 2004 to 2013. We studied the spatial‐temporal changes in movement behavior of 11 elephants in their core areas. Past studies have focused on elephant movement along corridors. We tested for the effect of poaching risk on their path straightness (i.e., tortuosity) while controlling for other environmental and human activities in the landscape using a set of generalized linear mixed models. To test for temporal variation of tortuosity, we used a time‐series linear model. Elephants turned less frequently while they were in poaching locations and at times with a high level of poaching activity, even though their speed did not change. The variation of tortuosity is a good indicator of differences in poaching risk as perceived by the elephants, which could complement patrol‐based anti‐poaching efforts by wildlife managers, especially in remote, inaccessible landscapes.

Connectivity within populations affects important ecological and evolutionary processes like gene flow, disease dynamics, and cultural exchange (Noad et al., 2000; McCallum & Dobson, 2002; Epps et al., 2005). Understanding connectivity is especially relevant in the context of conservation as landscape-level changes may alter wildlife movement. Such changes disproportionately affect wideranging species (Seidler et al., 2015), and those whose movement corridors are not protected (Didier et al., 2011). However, assessing connectivity across and within populations is difficult given the ephemeral and often cryptic nature of dispersal. Comparative, long-term data sets of known individuals can provide critical information and insights for wildlife managers and policymakers to determine whether and how subpopulations are connected. The Laikipia–Samburu elephant (Loxodonta africana Blumenbach) population is the second largest in Kenya with approximately 7415 individuals, primarily relying on range outside of governmentally protected areas (Poole et al., 1992; Litoroh et al., 2010). The 34,000 km2 Laikipia–Samburu ecosystem is a complex land use mosaic comprised of private, government and community lands (Thouless, 1995), which represent varying levels of risk to the region’s elephants (Ihwagi et al., 2015). The region is undergoing large-scale development projects (LAPSSET)

Social environments are fundamental to fitness in many species. In disrupted societies, the loss of important partners may alter social environments for surviving individuals. African elephants, Loxodonta africana, have experienced age-selective mortality linked to the ivory trade, and the resulting social costs for surviving young elephants are unknown. In this study, we followed orphaned female elephants and nonorphaned counterparts in Kenya's Samburu and Buffalo Springs National Reserves to elucidate whether orphaning and related dispersal behaviour incur social costs. There were clear social differences between orphans and nonorphans, most notably in that orphans tended to receive more aggression than nonorphans. Dispersal from natal groups was a behaviour found exclusively among orphans. Differences in social environments of orphans that remained in their natal groups and those that dispersed were also found in the form of dispersed orphans receiving more aggression while feeding than those that remained in their natal group. Our results suggest that orphaning in elephants is associated with social costs, and that these costs are amplified for orphans that disperse from their natal groups. Future research should identify the relationship between the social costs of being an orphan and fitness, which may be important to the recovery of populations affected by the ivory trade and other forms of disruption.

ABSTRACT Honeybees are globally recognized for products such as honey and wax, and as valuable pollinators of both natural ecosystems and agricultural crops. However, studies have shown that climate variability and human - driven environmental changes are affecting the population dynamics of the bees and their preferred fodder plants and subsequently, the socio-economic benefits of the honey bees. Although the decline in the honey bee and their associated plants may be attributed to all these factors combined, which rarely acts in isolation, previous studies in honey bee pollinator and pollination interactions have rarely considered they together decline. This study therefore aimed to investigate the effects of the combined and interactive factors of climate and anthropogenic environmental change on the bee forage diversity, plant-honey bee pollinators’ interactions and bee keeping activity. The project interviewed 25 respondents of smallholder farmer/beekeeper households using a semi-structured questionnaire. The beekeepers and other key informants in the study site were asked to report on important constrains and opportunities for beekeeping. Transect walks were conducted during the wet and dry season to determine the diversity of bee forage plants in twelve (12) randomly selected farms lined while bee population was determined by observing the beehive fences. Rainfall data collected in the same period were subsequently built into a statistical model to predict relationship between diversity of bee forage plants and bee population using precipitation data. Descriptive statistics were used to analyze quantitative data and percentages for the qualitative data. The major findings of the study indicated that there was a positive relationship between the warmer and drier weather conditions experienced during dry season and the lower diversity of bee forage plants [Bee forage plants (FP) (at confidence range) = 12.425 + 0.8757M, p-value; R2 = 0.8]. Similarly, there was also a positive correlation between the honey bee (B) population size and the availability of bee forage plants [(FB) = 17.116 + 0.6365 P, R2 = 0.55]. The findings indicated that warmer and drier conditions in dry season were accompanied with about 57% decline in the diversity of the honey bee fodder (floral resources) and about 36% decline in honeybee population. The most important plant families observed to be used by honey bee as fodder included Acanthaceae, Labiatae, Rubiaceae and Compositae. Among the plant species in the understory community, Tridax procumbens, Digera muricata and Justicia flava were found to be among the most important to honey bees. Hence, this study show clear evidence of a link between climate variability, diversity of honey bee fodder plants and honey bee population. The findings of this study recommends that beekeeping farmers in the study site should give consideration to the season long fodder resources needed by bees in dry season and ensure connectivity of natural habitats in farming areas, so that bees’ can more easily disperse and easily collect floral resources essential in response to changing climates.

Abstract As human and elephant populations grow in Kenya so does human-elephant conflict. One of the most substantial contributors to this conflict, the crop-raiding behavior of elephants (Loxodonta africana), is alleviated through the use of The Elephants and Bee Project's beehive fences. A threat to these beehives are the honey badgers (Mellivora capensis) who try to obtain honey, causing damage to the hive and the colony to abscond. The objective of this study was to improve the effectiveness of these beehive fences through identifying and testing novel honey badger deterrent methods. On-farm experiments in Taita Taveta County, Kenya were conducted to determine if visual and tactile deterrents could reduce the frequency and severity of honey badger hive predation of the hives compared to a previously used method. Prior to the start of the study, 77.1% percent of hives absconded within a week following a honey badger attack. After the addition of the novel deterrents (motion activated light deterrent, cone baffle, and hive cage deterrent), only 11.1% percent of the hives attacked by honey badgers absconded, suggesting the deterrents effectively reduced the amount of successful honey badger attacks. No relationship was found between deterrent type and amount of damage, nor for the duration and deterrent type. All deterrent methods effectively prevented honey badgers from raiding hives with variance in the success rates and economic feasibility. This project complemented the Elephants and Bee Project's ongoing research by providing much-needed insight into the role honey badger deterrents could play in preventing damage to the elephant deterring beehive fences. This research purposes deterrent recommendations based on cost effectiveness and ability to reduce honey badger raiding. These deterrents not only reduce honey badger hive raiding but also to improve human-honey badger coexistence as well as human-elephant coexistence.

The illegal killing of elephants, i.e. poaching and human-elephant related mortality, is the greatest immediate threats to elephants. They have led to declining of many populations of elephants in Africa. The Monitoring of Illegal Killing of Elephants (MIKE) program of the Convention on International Trade in Endangered Species (CITES) was set up in the year 2002 as a framework of monitoring trends in illegal killing in 57 African sites. MIKE program seeks to establish the relationships between the levels of illegal killing of elephants and various possible explanatory variables within and beyond the monitoring sites. The effort in implementing MIKE program vary from site to site, and to make the results comparable; a metric referred to as the Proportion of Illegally Killed Elephants (PIKE) out of all recorded deaths in a site has been adopted as the standard measure of severity of illegal killing. Loss of habitat due to the expansion of agriculture and infrastructural developments are the largest long-term threats to elephants. The migratory corridors of elephants and other wildlife in many landscapes have been cut off. The majority of wildlife resides outside formally protected areas on private and community lands. In the landscapes shared by wildlife and humans, competition for resources influences the spatial-temporal distributions of wildlife. Efforts to win the goodwill of private and community landowners regarding hosting of wildlife on their lands are ongoing in many sites across the elephant range. Despite the numerous studies on the nature of risk faced by elephants, fewer studies have focused on the behavioural adaptations of elephants living in those risky landscapes. This thesis sought to understand the site level drivers of illegal killing and how elephants adapt to the threat in Africa’s most intensively monitored site, the Laikipia-Samburu MIKE in northern Kenya. Using field verified records of causes of elephant mortality, the distribution of live elephants, and, the cadastral attributes of land parcels in the ecosystem, the thesis established that land use type is the most important correlate of levels of illegal killing and not its ownership. The study analyses the movement of elephants at hourly, day and night, and overall 24 hr activity cycle in relation to the spatial and temporal variation of the levels of illegal killing. Past studies have given a lot of attention to movement behaviour along corridors. The research in this thesis focusses on movement within core areas. At the hourly time interval, the research showed that elephants walk with lower tortuosity when they are in core areas with higher levels of illegal killing, i.e., higher risk. The study found that elephants move more at night when they are in core areas with higher risk, than when they are in safer core areas. Based on this finding, the research presents a new metric for inferring the levels of risk, i.e., night-day sped ratio. When elephants move from a core area to another one with a different level of risk, they alter their daily activity pattern to include a longer resting phase during the mid-day hours, and this is even more pronounced in core areas closest to permanent human settlements. The study found that as a result of the alteration of activity cycle within 24-hour periods, elephants loose approximately one hour of activity time. The results have the potential use as a remote means of assessing the spatial and temporal variation of risk by analysing elephant movement behaviour remotely thus complimenting patrol based anti-poaching efforts. The study provides new insight into the ecology of elephants living in fear. The confirmed increase of night-time movement potentially predisposes calves to the savannah predators, who are more active at night.

Projects updates from Save The Elephants

ABSTRACT As human and elephant populations grow in Kenya, elephants increasingly leave parks to eat farmers’ crops while foraging, which creates epicenters of human-elephant conflict (HEC). This conflict compromises farmers’ food and economic security, impedes elephant conservation initiatives, and threatens the safety of humans and elephants. In recent years, the situation has been exacerbated by drought and national-level infrastructure development that bisects key elephant habitat. Although researchers have widely studied elephant populations, few have examined the cultural, economic, and emotional effects of HEC on subsistence farmers. This project utilized a mixed methods approach to address this knowledge gap and understand the lived experiences of Wasaghala farmers in Lower Sagalla, Kenya. These farmers live adjacent to Kenya’s largest elephant population in Tsavo East National Park and regularly experience elephant crop-raiding. This research was conducted in partnership with Save the Elephants, a non-profit that studies elephant-crop raiding in Lower Sagalla. This project complements their research by facilitating greater understanding of complex human-elephant interactions and providing insight into the role that agricultural crops play in elephant crop-raiding. Personal interviews were conducted with a purposefully chosen sample of farmers, community leaders, and regional experts to understand their perspectives on cultural, agricultural, and economic dimensions of HEC in Lower Sagalla. Topics covered included regional history of HEC, impacts on farmers, elephant deterrent strategies, and farmer agricultural decision-making. Additional data were collected from an on-farm experiment that examined how crop palatability impacts elephant crop-raiding behavior. It aimed to determine if moringa and sunflowers are less palatable to elephants than maize, and if growing these crops can reduce crop loss due to elephant crop-raiding. Results from all data concluded that HEC creates widespread suffering for farmers in Lower Sagalla, that they are unable to adequately address this issue on their own, and that there is a need for the development of novel HEC mitigation strategies. Additionally, results suggest that crop palatability influences elephant crop-raiding behavior and that growing crops that are less palatable to elephants, but beneficial to farmers, may play a role in reducing crop loss and increasing farmers’ economic and food security. The research concludes with management recommendations to reduce elephant crop-raiding and improve human-elephant coexistence.

Seismic waves — vibrations within and along the Earth’s surface — are ubiquitous sources of information. During propagation, physical factors can obscure information transfer via vibrations and influence propagation range [1]. Here, we explore how terrain type and background seismic noise influence the propagation of seismic vibrations generated by African elephants. In Kenya, we recorded the ground-based vibrations of different wild elephant behaviours, such as locomotion and infrasonic vocalisations [2], as well as natural and anthropogenic seismic noise. We employed techniques from seismology to transform the geophone recordings into source functions — the time-varying seismic signature generated at the source. We used computer modelling to constrain the propagation ranges of elephant seismic vibrations for different terrains and noise levels. Behaviours that generate a high force on a sandy terrain with low noise propagate the furthest, over the kilometre scale. Our modelling also predicts that specific elephant behaviours can be distinguished and monitored over a range of propagation distances and noise levels. We conclude that seismic cues have considerable potential for both behavioural classification and remote monitoring of wildlife. In particular, classifying the seismic signatures of specific behaviours of large mammals remotely in real time, such as elephant running, could inform on poaching threats.

Repeated use of the same areas may benefit animals as they exploit familiar sites, leading to consistent home ranges over time that can span generations. Changing risk landscapes may reduce benefits associated with home range fidelity, however, and philopatric animals may alter movement in response to new pressures. Despite the importance of range changes to ecological and evolutionary processes, little tracking data have been collected over the long-term nor has range change been recorded in response to human pressures across generations. Here, we investigate the relationships between ecological, demographic and human variables and elephant ranging behaviour across generations using 16 years of tracking data from nine distinct female social groups in a population of elephants in northern Kenya that was heavily affected by ivory poaching during the latter half of the study. Nearly all groups—including those that did not experience loss of mature adults—exhibited a shift north over time, apparently in response to increased poaching in the southern extent of the study area. However, loss of mature adults appeared to be the primary indicator of range shifts and expansions, as generational turnover was a significant predictor of range size increases and range centroid shifts. Range expansions and northward shifts were associated with higher primary productivity and lower poached carcass densities, while westward shifts exhibited a trend to areas with higher values of primary productivity and higher poached carcass densities relative to former ranges. Together these results suggest a trade-off between resource access, mobility and safety. We discuss the relevance of these results to elephant conservation efforts and directions meriting further exploration in this disrupted society of a keystone species.

Abstract 1. Development of transportation corridors has accelerated globally, with infrastructure projects being implemented across remote ecosystems, particularly in the tropics. Such developments can have negative impacts on wildlife and their ecosystems. The importance of wildlife crossing structures to mitigate adverse effects of such features is widely recognized, but the siting of and investment in crossing structures is contentious. Data on animal movement provide valuable, highly specific information for such processes, but can present analytical challenges and remain underutilized in planning mitigation efforts. 2. We develop two algorithms based on Integer Linear Programming to prioritize crossing points based on frequency of use or breadth of coverage among tracked individuals. These scenarios represent metrics likely to guide the planning of crossing structures, where the former may relate to the objective of minimizing vehicle‐animal collisions and the latter on maintaining ecosystem connectivity. We exemplify the algorithms through application on a tracking dataset from over 150 African elephants living near the proposed Lamu Port‐South Sudan‐Ethiopia‐Transport corridor. We explore the influence of sampling bias on outcomes and discuss considerations to guide the application process. 3. Given the generally open, unfenced nature of this ecosystem, recorded movements occurred throughout the system and a third of the corridor length in the ecosystem was intersected by recorded elephant movements. The selection of crossing structure locations and their impacts on elephants varied whether we used a subsample of elephant representative of local population density or total sample of monitored individuals. The two algorithms also selected for different crossing structure locations. 4. Synthesis and applications. Our work shows some of the challenges of using Global Positioning System telemetry in deciding where to put crossing structures and demonstrates the need to identify the type of constraints in the system and desired crossing structure characteristics a priori. We recommend managers carefully evaluate the presence of potential biases in their data. High‐resolution data combined with objective prioritization methods allow reasoned planning actions, but are often lacking during critical infrastructure planning stages. Given the limited budget already allocated to mitigation measures in most proposed developments, the tools developed and applied here can facilitate effective spatial planning.

Abstract Network (graph) theory is a popular analytical framework to characterize the structure and dynamics among discrete objects and is particularly effective at identifying critical hubs and patterns of connectivity. The identification of such attributes is a fundamental objective of animal movement research, yet network theory has rarely been applied directly to animal relocation data. We develop an approach that allows the analysis of movement data using network theory by defining occupied pixels as nodes and connection among these pixels as edges. We first quantify node-level (local) metrics and graph-level (system) metrics on simulated movement trajectories to assess the ability of these metrics to pull out known properties in movement paths. We then apply our framework to empirical data from African elephants (Loxodonta africana), giant Galapagos tortoises (Chelonoidis spp.), and mule deer (Odocoileous hemionus). Our results indicate that certain node-level metrics, namely degree, weight, and betweenness, perform well in capturing local patterns of space use, such as the definition of core areas and paths used for inter-patch movement. These metrics were generally applicable across data sets, indicating their robustness to assumptions structuring analysis or strategies of movement. Other metrics capture local patterns effectively, but were sensitive to specified graph properties, indicating case specific applications. Our analysis indicates that graph-level metrics are unlikely to outperform other approaches for the categorization of general movement strategies (central place foraging, migration, nomadism). By identifying critical nodes, our approach provides a robust quantitative framework to identify local properties of space use that can be used to evaluate the effect of the loss of specific nodes on range wide connectivity. Our network approach is intuitive, and can be implemented across imperfectly sampled or large-scale data sets efficiently, providing a framework for conservationists to analyze movement data. Functions created for the analyses are available within the R package moveNT.

Living in Harmony with Elephants (LIHWE) is a four-part manual developed by Save the Elephants in collaboaration with Disney’s Animal Kingdom that teaches students conservation education with the aims to future wildlife ambassadors. Since it’s inception in 2012, LIHWE has reached out to over 500 students within the Samburu Isiolo Conservation Area (SICA). LIHWE is one of Save The Elephants (STE) Education Program’s approaches that introduce young minds into the world of elephants, delighting them in the species’ intelligence and diversity, and we remain indebted to Disney’s Animal Kingdom educators for all the help they provided to develop this manual. The first lesson equips students with knowledge on elephant ecology, exploring the species physiology, social interaction and its role in modifying the savannah. Next, learners are introduced to the importance of and threats to elephant habitats. The third lesson teaches learners how to protect themselves, their livestock, manyattas from elephants and while at it, securing a future for elephants and other wildlife.

The present thesis investigates aspects of the reproductive strategy of male African savannah elephants (Loxodonata africana). The existence of, and differences between alternative conditional dependent reproductive tactics are evaluated using a combination of behavioural, endocrinological and GPS tracking data and the age and tactic related success is measured using genetic paternity analysis. Hidden Markov Models were used as a probabilistic framework for analysing temporal changes in reproductively active and inactive periods based on shifts in association preferences of individuals. Distinct shifts between active and inactive periods were evident well before the onset of the aggressive reproductive tactic of musth, seen in older dominant males, hence providing the first quantitative evidence for the previously suggested sexually active periods in non-musth males. The link between hormones and reproductive status and tactics were investigated using a new technique for non-invasive faecal analysis of hormones. A combined analysis of androgens (Epiandrosterone) and glucocorticoid (3a,11-oxo-CM) hormones in relation to age, reproductive state and musth signals confirmed previously reported elevated levels of androgens during periods with temporal gland secretion and urine dribbling (Musth) but further showed that this increase is indeed linked to the presence of musth signals and not to the age of the individual. Androgen levels were generally increased during sexually active periods with a two-fold increase seen in active non-musth bulls and a four to six-fold increase in musth bulls. Contrary to expectations, increased glucocorticoids outputs were not seen in musth bulls but slightly elevated outputs were seen in active non-musth bulls. Brief peaks in glucocorticoids occurred immediately after physical fights and during periods of injuries. A general elevation of glucocorticoids was seen towards the end of the long dry season, a likely effect of ecological conditions. Behavioural changes and onset of musth signals occurred after an initial change in androgen levels suggesting that sex steroids may play an activiational role of sexually active periods as well as activation of the musth tactic within sexually active periods. Some evidence was found for increased androgen levels following encounters with receptive females, suggesting that such encounters may act as a “boosters vaccine” on androgen levels and hereby prolong active periods at times with high numbers of receptive females. Both musth and sexually active non-musth (SAN) bulls reduced their foraging and walked more than sexually inactive bulls. However musth bulls had a higher time specific investment in reproduction compared to SAN bulls with musth bulls doubling their daily walking distance compared to a 50% increase in SAN bulls. The (younger) non-musth bulls spent a larger part of the year sexually active (70%) compared to musth bulls (20%) hence SAN bulls have a tactic of prolonged low investment compared to the short and high investment seen in musth bulls, emphasising the need for evaluating the duration of reproductive bouts when comparing overall investments between tactics. The youngest (non-musth) bull to sire offspring was 21 years but older musth bulls above 35 years had a much higher age specific reproductive success compared to bulls below 30 years. However on a population level, bulls less than 30 years contributed 30% of the reproduction and 20-25% could be attributed to non-musth bulls.

Compensatory social behavior in nonhuman animals following maternal loss has been documented, but understanding of how orphans allocate bonding to reconstruct their social networks is limited. Successful social integration may be critical to survival and reproduction for highly social species and, therefore, may be tied to population persistence. We examined the social partners involved in affiliative interactions of female orphans and non-orphans in an elephant population in Samburu, northern Kenya that experienced heightened adult mortality driven by drought and intense ivory poaching. We contrasted partners across different competitive contexts to gain insight to the influence of resource availability on social interactions. Though the number of partners did not differ between orphans and non-orphans, their types of social partners did. Orphans interacted with sisters and matriarchs less while feeding than did non-orphans, but otherwise their affiliates were similar. While resting under spatially concentrated shade, orphans had markedly less access to mature adults but affiliated instead with sisters, bulls, and age mates. Orphan propensity to strengthen bonds with non-dominant animals appears to offer routes to social integration following maternal loss, but lack of interaction with adult females suggests orphans may experience decreased resource access and associated fitness costs in this matriarchal society.

Animal movement is fundamental for ecosystem functioning and species survival, yet the effects of the anthropogenic footprint on animal movements have not been estimated across species. Using a unique GPS-tracking database of 803 individuals across 57 species, we found that movements of mammals in areas with a comparatively high human footprint were on average one-half to one-third the extent of their movements in areas with a low human footprint. We attribute this reduction to behavioral changes of individual animals and to the exclusion of species with long-range movements from areas with higher human impact. Global loss of vagility alters a key ecological trait of animals that affects not only population persistence but also ecosystem processes such as predator-prey interactions, nutrient cycling, and disease transmission.

Introduction and background This joint report by Save the Elephants and Kenya Wildlife Service summarizes preliminary findings of a long-term study on the effects of the Standard Gauge Railway (SGR), the Mombasa - Nairobi highway, the Voi - Taveta highway and other infrastructural developments on elephant movements and ecosystem connectivity in the Tsavo ecosystem. The extent of the utilization of the wildlife crossing structures, examples of exceptional elephant movements, the effect of water distribution on elephant movements, the effects of a recently constructed fence-line along the SGR are assessed and reported on here. The report concludes by listing several management recommendations. The study began in March 2016 when 10 elephants were fitted with satellite tracking collars for monitoring their movements in relation to these new infrastructure projects.

Executive Summary From 2013 to 2016, Laos’s retail ivory market has expanded more rapidly than in any other country surveyed recently. ■ Laos has not been conforming with CITES regulations that prohibit the import and export of ivory. Since joining CITES in 2004, only one ivory seizure into Laos has been reported to the Elephant Trade Information System (ETIS). ■ Almost no arrests, let alone prosecutions and punishments, have been made of smugglers with ivory coming in or out of the country. ■ Most worked ivory for sale in Laos originates from elephants poached in Africa. ■ Ivory has also been entering Laos illegally from Thailand, as Thai traders have been offloading their ivory following the imposition of much stricter regulations there. ■ In late 2013 the average wholesale price of raw ivory sold by Lao traders peaked at about USD 2,000/kg. ■ By late 2016, the average wholesale price of raw ivory in Laos had declined to USD 714/kg, in line with prices elsewhere in the region. This price was much higher than in African countries, such as Sudan (Omdurman/Khartoum), where the average wholesale price of ivory was USD 279/kg in early 2017. This price differential is due to the extra expenses incurred in transport and bribes to government officials on the long journey to Asia. ■ In Laos, the decline in the wholesale price of raw ivory between 2013 and 2016, as elsewhere in the region, was mainly due to the slowdown in China’s economy, that resulted in an oversupply of illegal ivory, relative to demand. ■ Ivory items seen for sale in Laos are carved or machine-processed in Vietnam by Vietnamese and smuggled into Laos for sale, or are processed by Chinese traders in Laos on new computer driven machines. Ivory carving by Lao people is insignificant. ■ In Laos, the survey found 81 retail outlets with ivory on view for retail sale, 40 of which were in the capital, Vientiane, 21 in Luang Prabang, 8 in Kings Romans, 5 in Oudom Xay, 3 in Pakse, 2 in Dansavanh Nam Ngum Resort and 2 in Luang Nam Tha.

Poaching has escalated in recent years and is becoming the greatest immediate threat to elephants' survival. There is an urgent need to develop innovative and cost-effective methods for monitoring changes in elephant poaching levels remotely to complement the existing traditional field-based ground surveys. Since elephants are known to respond to anthropogenic risks by alterations in their speed of travel, we quantified this alteration as a ratio of night time speed to the day time speed (night-day speed ratio) and examined its relationship with poaching levels. Our hypothesis here is that poaching is a clear daytime risk, and thus an increase in night time movement rates over those seen during the day will support this hypothesis. Using elephant GPS tracking and mortality data collected in the Laikipia-Samburu ecosystem of northern Kenya between 2002 and 2012, we calculated the mean night-day speed ratio for collared elephants that utilised any of 13 contiguous land units, each under different ownership and management status, and related this ratio to the corresponding poaching levels before and during a poaching surge. Our study shows that the mean night-day speed ratio of both male and female elephants did not vary significantly by month, ruling out possible seasonal effect. However, both male and female elephants moved more at night than during the day where and when poaching levels were high. The relationship between poaching levels and night-day speed ratios was stronger for females than for males. We concluded that the variation in the night-day speed ratio of elephants might be used as an effective indicator for changes poaching levels on a near real-time basis. We recommend its adoption as a complimentary anti-poaching tool, where GPS tracking data is already available, because it would increase the geographical range for monitoring of poaching levels. The significant alteration in movement behaviour by elephants in response to poaching also has potential implications for their foraging strategy, reproduction and ultimate survival, all of which are not yet fully understood.

Kenya is endowed with an extraordinary wealth of mammals, birds and other biodiversity, a unique heritage for the people of Kenya. Kenya’s development blueprints on ensuring environmental sustainability – the Constitution (2010), Vision 2030, and Sustainable Development Goals (SDG) – recognize the importance of sustainable resource use, reducing biodiversity loss, and maintenance of ecosystems processes. In the Vision 2030, under the conservation strategic thrust, the flagship project on securing wildlife dispersal areas and migratory corridors features prominently as one of the economic and social pillars. All over the country, wildlife populations have declined dramatically over the last few decades. Ecosystems are failing to provide ecological services such as water storage, soil protection and climate moderation. At the same time, human-wildife conflict has increased. To reduce and reverse this trend, it is urgent to assess and secure Kenya’s wildlife dispersal areas and migratory corridors as a way to restore balance to our country’s superb environment.

The new Standard Gauge Railway (SGR) linking Mombasa to Nairobi became officially operational in June 2017. It is a flagship project for Kenya under Kenya’s blue print Vision 2030, whose goal is to transform Kenya into a middle-income industrialized economy by 2030. More than a quarter i.e., 135 km of the 487 km railway is through the Tsavo Conservation Area, bisecting the range of Kenya’s largest surviving single elephant population of 12,800 animals, as well as many other wildlife species. This presents a challenge. While the old railway line lay level with the ground, the new SGR is elevated up to 10 metres in some sections and fenced on either side, creating a substantial barrier to wildlife movement with likely negative consequences. The contractor of the SGR, the China Roads and Bridges Cooperation, built six official wildlife passages to connect Tsavo East to Tsavo West National Parks and Tsavo East to the Taita Ranches to allow animals to travel in search of food, water and mates. The 2 km Tsavo River super bridge, the Kenani and the Maungu railway crossing bridges makes a total of nine wildlife passages. Save the Elephants in partnership with the Kenya Wildlife Service has been tracking elephants to understand the effectiveness of these passages. We fitted ten elephants (eight along the SGR and two along the Voi – Taveta road) with GPS satellite radio transmitters in March 2016. In June 2016, we initiated systematic vehicle and foot monitoring along the SGR of elephants and other species, not fitted with radio transmitters, by using their signs such as footprints and dung. Some elephants have used them effortlessly, with families in tow, while others have preferred to use the culverts and bridges that perforate the line but which have not officially been classed as wildlife passages. Even though the details of other wildlife species crossings are not reported here, it is important to highlight here that giraffes generally avoided any form of underpasses with only one footprint recorded on 13th March 2017 at culvert DK234+062. Generally, however the culverts offer an opportunity for wildlife crossing points between the two sides of Tsavo National Park and adjoining ranches keeping genetic diversity open and a flow of inter-seasonal movements.

Dominance hierarchies are expected to form in response to socioecological pressures and competitive regimes. We assess dominance relationships among free-ranging female Asian elephants (Elephas maximus) and compare them with those of African savannah elephants (Loxodonta africana), which are known to exhibit age-based dominance hierarchies. Both species are generalist herbivores, however, the Asian population occupies a more productive and climatically stable environment relative to that of the African savannah population. We expected this would lower competition relative to the African taxon, relaxing the need for hierarchy. We tested whether 1) observed dominance interactions among individuals were transitive, 2) outcomes were structured either by age or by social unit according to 4 independent ranking methods, and 3) hierarchy steepness among classes was significant using David’s score. Elephas maximus displayed less than a third the number of dominance interactions as observed in L. africana, with statistically insignificant transitivity among individuals. There was weak but significant order as well as steepness among age-classes but no clear order among social units. Loxodonta africana showed significant transitivity among individuals, with significant order and steepness among age-classes and social units. Elephas maximus had a greater proportion of age-reversed dominance outcomes than L. africana. When dominance hierarchies are weak and nonlinear, signals of dominance may have other functions, such as maintaining social exclusivity. We propose that resource dynamics reinforce differences via influence on fission–fusion processes, which we term “ecological release.” We discuss implications of these findings for conservation and management when animals are spatially constrained.

Monitoring anthropogenic impacts on wildlife can be challenging, particularly when human activities affecting wildlife are cryptic. Using anti-predator behaviors as proxies for perceived pressure is appealing because of the relative ease with which they can be recorded and the presumed relationship between the threat of interest and a predator stimulus. However, behaviors are plastic and affected by factors unrelated to human activity. Consequently, it is critical to assess the relationship between behavioral indicators and their context before interpretation. In this study we used a combination of behavior, movement and demography from a threatened population of African elephants in northern Kenya to determine whether reaction to research vehicles was indicative of poaching pressure. We used mixed-effects models predicting reaction of elephants to observer vehicle approaches in which we treated individuals as random effects and included ecological, anthropogenic, spatial, social and demographic predictor variables. Contrary to our hypothesis, recorded levels of reactive behavior did not increase with poaching levels in either a population-level dataset or a data subset of individuals whose spatial behavior was precisely known via radio-tracking. Rather, primary productivity positively predicted reactive behavior in both datasets. This relationship was heightened by the presence of musth males in the radio-collar dataset. Reactivity was not related to the time since entering the protected areas, but increased among groups that spent less time in the protected areas. Inter-individual differences were apparent, suggesting the importance of inherent differences (e.g. personality) across groups. In our study, elephants plagued by a severe human threat did not react defensively to humans in another context, suggesting nuanced discrimination of threats. Our study demonstrates the caution that should be taken in designing studies that use behavioral indices to represent threat and contributes to a growing body of literature employing behavioral indicators to monitor wildlife populations of conservation concern.

Despite increased awareness, China continues to be the world’s major concern in the consumption of illegal ivory. Media attention and support from international NGOs have drawn attention to the poaching crisis currently facing elephants in Africa; there has been growing involvement and dialogue with China from Western leaders and other prominent gures about this. President Obama of the US and Prince William of the UK both spoke in 2015 on Chinese television addressing fears of the ongoing elephant poaching to meet demand for ivory in China. Within China certain celebrities have been involved in campaigns with NGOs, drawing further attention to the elephant-poaching crisis. There has been less awareness about ivory from the woolly mammoth as its tusks are legal, being from an extinct species; many people in China and around the world remain ignorant about the extent of the use of mammoth ivory in China

Determining underlying demographic population processes is fundamental for the management and monitoring of wildlife species (Caughley & Sinclair 1994). Data on demographic processes (fecundity, survival, age of first reproduction, etc.) allow quantification of population trajectories, the identification of population sectors to which growth is most sensitive and determinations of the mechanisms driving population trends (Caswell 2001). In particular, when animal populations have suffered severe declines, demographic data are critical for evaluating what conservation measures might be effective and estimating time to recovery (Beissinger & Westphal 1998). Such information is fundamental to policy debates regarding the viability and benefits of species trade.

Carbon-14 measurements on 231 elephant ivory specimens from 14 large ivory seizures (?0.5 ton) made between 2002 and 2014 show that most ivory (ca. 90%) was derived from animals that had died less than 3 y before ivory was confiscated. This indicates that the assumption of recent elephant death for mortality estimates of African elephants is correct: Very little “old” ivory is included in large ivory shipments from Africa. We found only one specimen of the 231 analyzed to have a lag time longer than 6 y. Patterns of trade differ by regions: East African ivory, based on genetic assignments of geographic origin, has a much higher fraction of “rapid” transit than ivory originating in the Tridom region of Cameroon–Gabon–Congo. Carbon-14 is an important tool in understanding patterns of movement of illegal wildlife products.

Dominance hierarchies are expected to form in response to socioecological pressures and competitive regimes. We assess dominance relationships among free-ranging female Asian elephants (Elephas maximus) and compare them with those of African savannah elephants (Loxodonta africana), which are known to exhibit age-based dominance hierarchies. Both species are generalist herbivores, however, the Asian population occupies a more productive and climatically stable environment relative to that of the African savannah population. We expected this would lower competition relative to the African taxon, relaxing the need for hierarchy. We tested whether 1) observed dominance interactions among individuals were transitive, 2) outcomes were structured either by age or by social unit according to 4 independent ranking methods, and 3) hierarchy steepness among classes was significant using David’s score.