Barnes, R.F.W., Douglas-Hamilton, I.
The Numbers and Distribution Patterns of Large Mammals in the Ruaha- Rungwa Area of Southern Tanzania (1982)
Journal of Applied Ecology Vol. 19, No. 2 (Aug., 1982), pp. 411-425

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Describes aerial census of Ruaha National Park and adjacent areas including Rungwa and Kisigo Game Reserves. The 31,500 sq km census zone carries one of the largest elephant populations in Africa with an estimated 43,865 elephants with 24,625 estimated to occur within the 10, 200 sq km Park itself. Within the park, comparisons with earlier counts showed an apparent increase in elephant density of 8-10% per annum since 1965. The increase in elephant within the park itself probably results from the change in human distribution in the region and a period of higher rainfall.

Bastille-Rousseau G., Douglas-Hamilton I., Blake S., Northrup J.M., Wittemyer G.
Applying network theory to animal movements to identify properties of landscape space use (2018)
Ecol Appl. 2018 Apr;28(3):854-864. doi: 10.1002/eap.1697.

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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.

Bastille‐Rousseau, G., Wall, J., Douglas‐Hamilton, I., Wittemyer, G.
Optimizing the positioning of wildlife crossing structures using GPS telemetry (2018)
J Appl Ecol. 2018;55:2055–2063. https://doi.org/10.1111/1365-2664.13117 First published: 06 February 2018

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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.

Bastille‐Rousseau, G., Wittemyer, G.
Characterizing the landscape of movement to identify critical wildlife habitat and corridors (2020)
Conservation Biology, Volume 0, No. 0, 1–14 © 2020 Society for Conservation Biology DOI: 10.1111/cobi.13519

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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.

Berti, E., Davoli, M., Buitenwerf, R., Dyer, A., Hansen, O.L., Hirt, M., Svenning, J.C., Terlau, J.F., Brose, U. and Vollrath, F.
The r package enerscape: A general energy landscape framework for terrestrial movement ecology (2022)
Methods in Ecology and Evolution, 13(1), pp.60-67.

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Abstract 1. Ecological processes and biodiversity patterns are strongly affected by how animals move through the landscape. However, it remains challenging to predict animal movement and space use. Here we present our new r package enerscape to quantify and predict animal movement in real landscapes based on energy expenditure. 2. enerscape integrates a general locomotory model for terrestrial animals with GIS tools in order to map energy costs of movement in a given environment, resulting in energy landscapes that reflect how energy expenditures may shape habitat use. enerscape only requires topographic data (elevation) and the body mass of the studied animal. To illustrate the potential of enerscape, we analyse the energy landscape for the Marsican bear (Ursus arctos marsicanus) in a protected area in central Italy in order to identify least- cost paths and high- connectivity areas with low energy costs of travel. 3. enerscape allowed us to identify travel routes for the bear that minimize energy costs of movement and regions that have high landscape connectivity based on movement efficiency, highlighting potential corridors. It also identifies areas where high energy costs may prevent movement and dispersal, potentially exacerbating human– wildlife conflicts in the park. A major strength of enerscape is that it requires only widely available topographic and body size data. As such, enerscape permits a first cost- effective way to estimate landscape use and movement corridors even when telemetry data are not readily available, such as for the example with the bear.

Blake S., Deem S.L., Strindberg S., Maisels, F., Momont, L., Isia, I., Douglas-Hamilton, I., Karesh, W.B., Kock, M.D.,
Roadless Wilderness Area Determines Forest Elephant Movements in the Congo Basin. (2008)
PLoS ONE 3(10): e3546. doi:10.1371/journal.pone.0003546

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A dramatic expansion of road building is underway in the Congo Basin fuelled by private enterprise, international aid, and government aspirations. Among the great wilderness areas on earth, the Congo Basin is outstanding for its high biodiversity, particularly mobile megafauna including forest elephants (Loxodonta africana cyclotis). The abundance of many mammal species in the Basin increases with distance from roads due to hunting pressure, but the impacts of road proliferation on the movements of individuals are unknown. We investigated the ranging behaviour of forest elephants in relation to roads and roadless wilderness by fitting GPS telemetry collars onto a sample of 28 forest elephants living in six priority conservation areas. We show that the size of roadless wilderness is a strong determinant of home range size in this species.

Blake, S.
The Ecology of Forest Elephant Distribution and its Implications for Conservation. (2002)
Ph.D. dissertation, University of Edinburgh, Edinburgh

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The 1980's saw African elephants massacred from an estimated continental population of 1.3 million in 1979 to just 609, 000 ten years later (Douglas-Hamilton 1989) killed mostly for their teeth. During that decade wildlife managers, conservationists, and some politicians battled to stem the slaughter in East and Southern Africa (Douglas-Hamilton and Douglas-Hamilton 1982, Douglas-Hamilton 1988, Cobb 1989 and Western 1989b) which culminated among other things in the ban of international trade in ivory (Sharp 1997). During this time of visible slaughter of savannah elephants (Loxodanta africana africana), there was a general feeling that the 'invisible' elephants (L. a. cyclotis) of the vast equitorial forests, largely uninhabited and unknown, were relatively free from poaching and that large numbers of elephants remained (Anon. 1984a; Owen-Smith 1988).

Blake, S., Bouché, P., Rasmussen, H., Orlando, A., Douglas-Hamilton, I.
Les derniers éléphants du Sahel. Comportement migratoire, état de la population et histoire récente des éléphants du désert du Mali. (2003)
Save The Elephants

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Les éléphants occupaient jadis un vaste territoire continu à travers toute l’Afrique de l’Ouest,?s’étendant des forêts côtières au Sahara. Le déclin de ces populations autrefois répandues, sous?l’effet du braconnage destiné au commerce de l’ivoire, de l’empiètement humain, et combinés à?l’absence de conservation et d’attention scientifique, se produit à un rythme alarmant. Les?populations qui subsistent sont de petite taille, extrêmement fragmentées et géographiquement?isolées, dont plus de la moitié ne contient plus aujourd’hui qu’une centaine d’individus à peine?(Roth and Douglas-Hamilton 1991 ; Said et al. 1995 ; Barnes et al. 1998 ; Barnes 1999).

Blake, S., Bouché, P., Rasmussen, H., Orlando, A., Douglas-Hamilton, I.
The Last Sahelian Elephants. Ranging Behavior, Population Status and Recent History of the Desert Elephants of Mali. (2003)
Save The Elephants

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Elephants once occupied a largely continuous range across West Africa, from the coastal forests to the Sahara. The collapse of these once extensive populations, caused by poaching for the ivory trade, human encroachment and the concurrent lack of conservation and scientific attention, has been alarming. Remaining populations are small, highly fragmented and geographically isolated, with over half now containing fewer than 100 individuals (Roth and Douglas-Hamilton 1991; Said et al. 1995; Barnes et al. 1998; Barnes 1999). The population living in the Gourma, which before this survey was estimated to be between 300 and 800, is one of the most important in the West African region and is accorded a high priority in the regional elephant strategy of the IUCN (Worldwide Conservation Union).

Blake, S., Douglas-Hamilton, I., Karesh W.
GPS telemetry of forest elephants in Central Africa: results of a preliminary study. (2001)
Afr. J. Ecol. 39(2), 178-186. doi.10.1046/j.1365-2028.2001.00296.x

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Few data exist on the ranging behaviour of forest elephants. A feasibility study on the use of GPS telemetry as a tool to study ranging, seasonal movements and distribution was implemented in the Dzanga-Sangha and Nouabale¨-Ndoki National Parks Complex of Central African Republic and Congo. The study consisted of two parts - a thorough hand-held testing of an elephant  GPS telemetry collar under tropical forest conditions and the deployment of collars on two elephants. During the feasibility study the system performance was satisfactory; GPS fix acquisition success rate, VHF and UHF collar-researcher communications were dequate. Two elephants, a mature bull and an adult female, were immobilized and fitted with GPS collars in October 1998. After deployment, the female's GPS collar performed well initially, but in less than a month the GPS within the collar stopped acquiring fixes.

Boettiger, A.N., Wittemyer, G., Starfield, R., Vollrath, F., Douglas-Hamilton, I., Getz, W.M.
Inferring ecological and behavioral drivers of African elephant movement using a linear filtering approach. (2011)
Ecology 92:1648–1657. [doi:10.1890/10-0106.1]

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Understanding the environmental factors influencing animal movements is fundamental to theoretical and applied research in the field of movement ecology. Studies relating fine-scale movement paths to spatiotemporally structured landscape data, such as vegetation productivity or human activity, are particularly lacking despite the obvious importance of such information to understanding drivers of animal movement. In part, this may be because few approaches provide the sophistication to characterize the complexity of movement behavior and relate it to diverse, varying environmental stimuli. We overcame this hurdle by applying, for the first time to an ecological question, a finite impulse–response signal-filtering approach to identify human and natural environmental drivers of movements of 13 free-ranging African elephants (Loxodonta africana) from distinct social groups collected over seven years.

Bohrer, G., Beck, P.S.A., Ngene, S.M., Skidmore, A.K., Douglas-Hamilton, I.
Elephant movement closely tracks precipitation-driven vegetation dynamics in a Kenyan forest-savanna landscape. (2014)
Movement Ecology 2014, 2:2 doi:10.1186/2051-3933-2-2 The electronic version of this article is the complete one and can be found online at: http://www.movementecologyjournal.com/content/2/1/2

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This study investigates the ranging behavior of elephants in relation to precipitation-driven dynamics of vegetation. Movement data were acquired for five bachelors and five female family herds during three years in the Marsabit protected area in Kenya and changes in vegetation were mapped using MODIS normalized difference vegetation index time series (NDVI). In the study area, elevations of 650 to 1100 m.a.s.l experience two growth periods per year, while above 1100 m.a.s.l. growth periods last a year or longer.

Bouché, P., Douglas-Hamilton, I., Wittemyer, G., Nianogo, A., Lejeune, P., Vermeulen, C.
Will Elephants Soon Disappear from West African Savannahs? (2011)
PLoS ONE | www.plosone.org 5 June 2011 | Volume 6 | Issue 6 | e20619

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Precipitous declines in Africa’s native fauna and flora are recognized, but few comprehensive records of these changes have been compiled. Here, we present population trends for African elephants in the 6,213,000 km2 Sudano-Sahelian range of West and Central Africa assessed through the analysis of aerial and ground surveys conducted over the past 4 decades. These surveys are focused on the best protected areas in the region, and therefore represent the best case scenario for the northern savanna elephants. A minimum of 7,745 elephants currently inhabit the entire region, representing a minimum decline of 50% from estimates four decades ago for these protected areas. Most of the historic range is now devoid of elephants and, therefore, was not surveyed. Of the 23 surveyed elephant populations, half are estimated to number less than 200 individuals.

Branco M.S, P.S., Merkle, J.A., Pringle, R.M., King, L., Tindall, T., Stalmans, M., Long, R.A.
An experimental test of community-based strategies for mitigating human–wildlife conflict around protected areas (2019)
Conservation Letters. 2019;e12679. wileyonlinelibrary.com/journal/conl 1 of 8 https://doi.org/10.1111/conl.12679

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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.

Burrill, A., Douglas-Hamilton, I.
African elephant database project: Final Report – Phase One. (1987)
WWF/GRID/UNEP

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The objective of this report is to provide information on elephant numbers, distribution and trends, and factors affecting these, which will be helpful to countries in reviewing and setting quotas under the CITES Ivory Export Quota agreement. ??The data on elephants have been acquired from published scientific literature, reports of aerial or ground surveys, and from a series of questionnaires distributed up to December 1986.

Butler, K., M.
Behaviour and crop-raiding patterns of Asian Elephants (Elephas maximus): Can beehive fences help mitigate human-elephant conflict in Sri Lanka? (2020)
B.Sc.(Ecology and Sustainability) (Hons.), M. Env A thesis submitted in total fulfillment of the requirements for the degree of Master of Philosophy (Environmental Science) Faculty of Science School of Environmental and Life Sciences University of Newcastle New South Wales, Australia August 2019

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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.

Canney, S., Lindsey, K., Hema, E., Douglas-Hamilton, I., Martin, V.
Initiative pour les éléphants du Mali: une synthèse des connaissances et des recherches, des recommandations relatives au niveau de population, à la surface de l’habitat et aux menaces qui pèsent sur les éléphants de Gourma. (2007)
Save the Elephants

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L’équipe du Projet Eléphant Mali constituée de la WILD Foundation, de Save the Elephants et de l’Environment and Development Group est heureuse de soumettre ce rapport sur la première phase (2003-2006) du projet. Le projet s’est appuyé sur les travaux effectués précédemment par Save the Elephants et d’autres, et consistait en des travaux originaux de recherche scientifique sur le terrain et la compilation d’informations d’archivage pouvant servir de base à un plan de gestion pour la survie des éléphants du Gourma. ?Ce projet a été rendu possible grâce à l’aide financière, technique et en nature de nombreuses agences, organisations et individus.?Le gouvernement du Mali – le projet présentait l’indéniable avantage de recevoir l’appui explicite du Président de la République du Mali, Alpha Oumar Konare et de son successeur, Amadou Toumani Toure.

Canney, S., Lindsey, K., Hema, E., Douglas-Hamilton, I., Martin, V.
The Mali elephant initiative: synthesis of knowledge, research and recommendations about the population, its range and the threats to the elephants of the Gourma. (2007)
Save the Elephants

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The Gourma elephant population is unique in Africa for three reasons: it is the northernmost population on the continent, it occupies an exceptionally harsh, arid environment and it owes its existence to historical co-existence with the people of the region. The people and government of Mali have much to be proud of, in their preservation of this valuable cultural and biological treasure. However, pressures on both people and elephants are growing, and accurate knowledge is essential for the development of well-informed strategies if this population of elephants is to have a long-term future.?The aim of the initiative was to better understand: • the current size, composition and status of the elephant population • the ecological requirements of the elephants? • patterns of human activity and their influence on the human-elephant relationship/elephant livelihoods

Cerling, T.E., Barnettea, J.E., Chessona, L.A., Douglas-Hamilton, I., Gobush, K.S., Unog, K.T., Wasser, S.K., Xui, X.
Radiocarbon dating of seized ivory confirms rapid decline in African elephant populations and provides insight into illegal trade (2016)
www.pnas.org/cgi/doi/10.1073/pnas.1614938113

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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.

Cerling, T.E., Wittemyer G., Ehleringer J.R., Remien C.H., Douglas-Hamilton, I.
History of Animals using Isotope Records (HAIR): A 6-year dietary history of one family of African elephants. ()
Proceedings of the National Academy of Sciences www.pnas.org/cgi/doi/10.1073/pnas.0902192106

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The dietary and movement history of individual animals can be studied using stable isotope records in animal tissues, providing insight into long-term ecological dynamics and a species niche. We provide a 6-year history of elephant diet by examining tail hair collected from 4 elephants in the same social family unit in northern Kenya. ??Sequential measurements of carbon, nitrogen, and hydrogen isotope rations in hair provide a weekly record of diet and water resources. Carbon isotope ratios were well correlated with satellite-based measurements of the normalized difference vegetation index (NDVI) of the region occupied by the elephants as recorded by the global positioning system (GPS) movement record; the absolute amount of C4 grass consumption is well correlated with the maximum value of NDVI during individual wet seasons.

Cerling, T.E., Wittemyer, G., Rasmussen, H.B., Vollrath, F., Cerling, C.E., Robinson, T., Douglas-Hamilton, I.
Stable isotopes in elephant hair document migration patterns and diet changes. (2006)
PNAS January 10, 2006 vol. 103 no. 2 371–373

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We use chronologies of stable isotopes measured from elephant (Loxodonta africana) hair to determine migration patterns and seasonal diet changes in elephants in and near Samburu National Reserve in northern Kenya. Stable carbon isotopes record diet changes, principally enabling differentiation between browse and tropical grasses, which use the C3 and C4 photosynthetic pathways, respectively; stable nitrogen isotopes record regional patterns related to aridity, offering insight into localized ranging behavior. Isotopically identified range shifts were corroborated by global positioning system radio tracking data of the studied individuals. Comparison of the stable isotope record in the hair of one migrant individual with that of a resident population shows important differences in feeding and ranging behavior over time.

Chan, A.N., Wittemyer, G., McEvoy, J., Williams, A.C., Cox, N., Soe, P., Grindley, M., Shwe, N.M., Chit, A.M., Oo, Z.M. and Leimgruber, P.
Landscape characteristics influence ranging behavior of Asian elephants at the human-wildlands interface in Myanmar (2022)
Movement Ecology, 10(1), pp.1-15.

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Context Asian elephant numbers are declining across much of their range driven largely by serious threats from land use change resulting in habitat loss and fragmentation. Myanmar, holding critical range for the species, is undergoing major developments due to recent sociopolitical changes. To effectively manage and conserve the remaining populations of endangered elephants in the country, it is crucial to understand their ranging behavior. Objectives Our objectives were to (1) estimate the sizes of dry, wet, and annual ranges of wild elephants in Myanmar; and quantify the relationship between dry season (the period when human-elephant interactions are the most likely to occur) range size and configurations of agriculture and natural vegetation within the range, and (2) evaluate how a percentage of agriculture within dry core range (50% AKDE range) of elephants relates to their daily distance traveled.

Chiyo, P. I., Grieneisen, L. E., Wittemyer, G., Moss, C.J., Lee, P. C., Douglas-Hamilton, I., Archie, E. A.
The influence of social structure, habitat, and host traits on the transmission of Escherichia coli in wild elephants. (2014)
PLoS One e93408. DOI: 10.1371/journal.pone.0093408. http://dx.doi.org/10.1371/journal.pone.0093408

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Social structure is proposed to influence the transmission of both directly and environmentally transmitted infectious agents. However in natural populations, many other factors also influence transmission, including variation in individual susceptibility and aspects of the environment that promote or inhibit exposure to infection. We used a population genetic approach to investigate the effects of social structure, environment, and host traits on the transmission of Escherichia coli infecting two populations of wild elephants: one in Amboseli National Park and another in Samburu National Reserve, Kenya. If E. coli transmission is strongly influenced by elephant social structure, E. coli infecting elephants from the same social group should be genetically more similar than E. coli sampled from members of different social groups. However, we found no support for this prediction.

Cook, R.M., Parrini, F., King, L.E., Witkowski, E.T.F., Henley, M.D
African honeybees as a mitigation method for elephant impact on trees (2017)
Elsevier. Biological Conservation Volume 217, January 2018, Pages 329-336 https://doi.org/10.1016/j.biocon.2017.11.024

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Conservation managers are concerned about the impact that African elephants (Loxodonta africana) have on large tree species, necessitating the need for mitigation methods. Elephants actively avoid contact with African honeybees (Apis mellifera subsp. scutellata), staying clear of crop fields surrounded by beehive fence-lines and moving away from the sounds of swarming honeybees. Therefore, our objectives were to test whether the presence of beehives in trees influenced the likelihood of the tree receiving elephant impact, and compare these results to wire-netted (method used to prevent bark-stripping) and control (no treatment) trees. We selected a tree highly sought after by elephant, the marula tree (Sclerocarya birrea subsp. caffra), as our study species. We also assessed whether elephants avoided areas with marula trees containing beehives. Finally we provide a comparison of the financial costs of the beehive and wire-netting mitigation methods. We hung 50 active beehives in 50 trees, with 50 dummy beehives hung from branches on the opposite ends of each tree's main stem. We wire-netted another 50 trees and then assigned 50 trees as a control. Elephant impact on all 150 trees was measured prior to the addition of treatments and then post-treatment addition for 9 months. 54% of the control trees received some form of elephant impact, in comparison to 28% of the wire-netted trees and only 2% of the beehive trees. Wire-netting protected trees against bark-stripping but did not prevent elephants from breaking branches. Beehives proved to be the more effective mitigation method for elephant impact on large trees, although the presence of beehives did not prevent elephants from moving through the study site. The financial cost and maintenance required for the beehive mitigation method are greater than that of wire-netting, but the beehives can provide honey as an additive benefit on a small-scale usage level.

De Knegt, H.J. et al
The spatial scaling of habitat selection by African elephants. (2010)
Journal of Animal Ecology. DOI: 10.1111/j.1365-2656.2010.01764.x

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1. Understanding and accurately predicting the spatial patterns of habitat use by organisms is?important for ecological research, biodiversity conservation and ecosystem management. However,?this understanding is complicated by the effects of spatial scale, because the scale of analysis?affects the quantification of species–environment relationships.?2. We therefore assessed the influence of environmental context (i.e. the characteristics of the landscape?surrounding a site), varied over a large range of scales (i.e. ambit radii around focal sites), on?the analysis and prediction of habitat selection by African elephants in Kruger National Park,?South Africa.

de Silva, S., Schmid, V., Wittemyer, G.
Fission–fusion processes weaken dominance networks of female Asian elephants in a productive habitat (2016)
Behavioral Ecology (2016), 00(00), 1–10. doi:10.1093/beheco/arw153

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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.

de Silva, S., Schmid, V., Wittemyer, G.
Fission–fusion processes weaken dominance networks of female Asian elephants in a productive habitat (2017)
Behavioral Ecology (2017), 28(1), 243–252. doi:10.1093/beheco/arw153

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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.

de Silva, S., Wittemyer, G.
A Comparison of Social Organization in Asian Elephants and African Savannah Elephants. (2012)
Int J Primatol. DOI 10.1007/s10764-011-9564-1

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Asian and African elephant species have diverged by ca. 6 million years, but as large, generalist herbivores they occupy similar niches in their respective environments. Although the multilevel, hierarchical nature of African savannah elephant societies is well established, it has been unclear whether Asian elephants behave similarly. Here we quantitatively compare the structure of both species’ societies using association data collected using the same protocol over similar time periods. Sociality in both species demonstrates well-defined structure, but in contrast to the African elephants of Samburu the Uda Walawe Asian elephants are found in smaller groups, do not maintain coherent core groups, demonstrate markedly less? ?social connectivity at the population level, and are socially less influenced by seasonal differences in ecological conditions.

Didier, K.A., Cotterill, A., Douglas-Hamilton, I., Frank, L., Georgiadis, N.J., Graham, M., Ihwagi, F., King, J., Malleret-King, D., Rubenstein, D., Wilkie, D., Woodroffe, R.
Landscape-Scale Conservation Planning of the Ewaso Nyiro: A Model for Land Use Planning in Kenya? (2006)
Landuse Planning Workshop Report, Mpala

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The unique wildlife of the Ewaso Nyiro and valuable services that the?ecosystem provides for humans (e.g., clean water and productive grasslands) cannot be?conserved by working solely on traditional conservation strongholds such as the national?reserves and private ranches of central Laikipia. To reach objectives for conserving wildlife,?stakeholders must work to preserve wildlife habitat and corridors in the surrounding?human-dominated landscape—a daunting task considering the complexity of working?at large spatial scales (e.g., many landowners, competing land uses) and limited conservation?resources available. Systematic, landscape-scale conservation planning helps?stakeholders set meaningful and transparent objectives, identify where to work to meet?those objectives, and prioritize areas for immediate investment.

Didier, K.A., Wilkie, D., Douglas-Hamilton, I., Frank, L., Georgiadis, N., Graham, M., Ihwagi, F., King, A., Cotterill, A., Rubenstein, D., Woodroffe, R.
Conservation Planning on a Budget: a “Resource Light” Method for Mapping Priorities at a Landscape Scale? (2009)
Biodivers Conserv (2009) 18:1979–2000 DOI 10.1007/s10531-008-9568-0

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Conservation projects may be reluctant to attempt Systematic Conservation Planning because existing methods are often prohibitive in the time, money, data, and expertise they require. We tried to develop a ‘‘resource light’’ method for Systematic Conservation Planning and applied it to the Ewaso Ngiro Landscape of central Kenya. Over a 6-month preparation period and 1-week participatory workshop, we used expert assessments to select focal biodiversity features, set quantitative targets for these, map their current distribution, vulnerability, potential for recovery, and conservation costs, and, finally, map cross-feature conservation priorities. Preparation for and facilitation of the workshop required time investment by one part-time workshop coordinator, eight workshop committee members, six ecosystem experts, and two GIS technicians.

Douglas-Hamilton, I.
Ivory an.d Insecurity: The Global Implications of Poaching in Africa (2012)
Testimony before the Committee on Foreign Relations U.S. Senate. US Government Printing Office.

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INTRODUCTION Mr. Chairman, Mr. Ranking Member, and members of the Committee, thank you for the opportunity to testify today. I am honoured to appear before your committee. My name is Iain Douglas-Hamilton, and I have been studying elephants in Africa since 1966. I founded the African Elephant Specialist Group of the IUCN in 1975 and launched the first pan-African elephant survey in 1977, funded by WCS and WWF. I have testified three times before Congress in the 1970s and 1980s when elephants were in peril from the ivory trade. We are once again in that situation. We are experiencing a huge upsurge in poaching, possibly to levels as high as those witnessed in the 1980s before the ivory ban. This time, however, we have more eyes on the ground and some unified systems endorsed by parties of the CITES treaty, for monitoring illegal killing of elephants (MIKE).

Douglas-Hamilton, I.
Another look at the ivory trade (Viewpoint) (1984)
Africa Wildlife Foundation News

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Refutes the hypothesis that the ivory trade does not pose an immediate crisis for the elephant, but that the increase in human population causing the restriction in elephant range is responsible for loss of elephants and is at the root of current population decline. Illustrates his criticism of the hypothesis by providing import and export figures. Suggests civil strife has played an important role in the loss of elephant population and the anarchy associated with this situation allows rampant poaching and hence illegal ivory trading to go on. Article could be considered a response to Parker and Amin's 'Ivory Crisis'.

Douglas-Hamilton, I.
Ivory price rises and inflation (1983)

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The paper relates ivory price to an index of inflation in order to explore the basis of a link between the rate of the killing of elephants and the rise in price of ivory greater than that of inflation. The index of inflation used came from the OECD for industrialized countries increased by 3.5 time between 1960 and 1980. Ivory prices for the similar period are taken from Parker (1979) and Parker and Bradley Martin (1983) using 1960 prices as the base year. Between 1960 and 1968 ivory prices kept broadly in line with the rate of inflation. Between 1969 and 1978 ivory prices exceeded inflation by a factor of 5.9 and between 1978 to 1982 by a factor of 3.5. The higher rate of inflation in individual elephant producing countries such as Kenya, Zaire and Uganda would it is argued reinforce the demand for ivory as a valuable commodity.

Douglas-Hamilton, I.
The current elephant poaching trend. (2009)
Pachyderm No. 45 July 2008–June 2009

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African and Asian elephants are in for tough times ahead. Their problems are complex. In southern Africa worries are still expressed about ‘too many elephants’ destroying woody vegetation in protected areas. In most of their range, crop raiding and conflict with human beings is reported every week. Currently, there are at least two horrendous droughts in Africa with negative effects on elephants-one in northern Kenya and another in Mali-perhaps related to widespread climate change but very much aggravated by habitat degradation caused by livestock overgrazing. Almost everywhere that elephants live, there is an ever-expanding, resource-hungry human population, with many who live hand- to-mouth. An enormous challenge lies ahead of reconciling conservation with poverty alleviation, and yet leaving adequate space for elephants and other wildlife.

Douglas-Hamilton, I.
Wilderness From an Elephant’s Point of View (2004) Wilderness and Human Communities. (2004)
Proceedings from the 7th World Wilderness Congress

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Wilderness and Human Communities. Proceedings from the 7th World Wilderness Congress 2004 The African wilderness, for me, isepitomized by elephants, so let us con­sider wilderness from an elephant's point of view. Elephants need a great deal of space. So from an elephant'spoint of view, the more wilderness the better. At present, where elephants dowell in Africa there is much habitat to support them, and so a host of otherspecies is surviving along with them. In this sense, ele­phants are anindicator of the welfare of the wilderness. Despite serious episodes of decrease over the last thirty years in their range north of the Zambezi, elephants still occur in abundance in huge and often remote wild land areas in Africa, and are still believed to have a total range of over 5 million square kilometers.

Douglas-Hamilton, I.
‘May he walk in peace’ – A tribute to PRINCE BERNHARD OF THE NETHERLANDS (1911 – 2004) (2004)
SWARA October – December 2004

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Iain Douglas-Hamilton remembers a man who fought long and hard to advance the conservation cause in Africa – and around the world. Prince Bernhard of the Netherlands died of cancer on 1 December 2004 at the age of 93. While he had a long and distinguished career in Europe’s turbulent 20th Century, he will be remembered in Africa primarily as one who tirelessly promoted the wildlife cause over many years. All his life he used a position of privilege to fight for matters of principle. As a German Prince he married Princess Juliana of the Dutch royal family. When Germany invaded his adopted country he backed the Free Dutch at a time when it looked as though the Nazis could never be defeated. He became a Dutchman among the Dutch, and a symbol of the resistance. And when the allies finally triumphed he never forgot the ordinary people who had joined the resistance.

Douglas-Hamilton, I., Bhalla, S., Wittemyer, G., Vollrath, F.
Behavioural reactions of elephants towards a dying and deceased matriarch (2006)
Applied Animal Behaviour doi:10.1016/j.applanim.2006.04.014

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The extent to which elephants hold behavioural traits in common with human beings is relevant to the ethics of how we treat them. Observations show that elephants, like humans, are concerned with distressed or deceased individuals, and render assistance to the ailing and show a special interest in dead bodies of their own kind. This paper reports helping and investigative behaviour of different elephants and their families towards a dying and deceased matriarch. We make use of long-term association records, GPS tracking data and direct observations. Records made around the time of death, shows that the helping behaviour and special interest exhibited was not restricted to closely related kin. The case is made that elephants, like human beings, can show compassionate behaviour to others in distress.

Douglas-Hamilton, I., Burill, A., Mackinnon, J.
Protected Areas as Refugees for Elephants. (1986)
Gland, IUCN

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Attempts to determine if vegetation types of Africa are adequately covered in the existing protected areas system. The elephant, which can exist within a very wide range of vegetation types, makes a particularly suitable indicator species because it is large and conspicious, and data on its distribution can be easily collected from both air and ground surveys. Results of distribution mapped from the 1979 African Elephant Survey and Conservation Programme and the protected areas were superimposed on White's vegetation map of Africa. The percentage of current elephant range and the total area for each vegetation zone within the biogeographical units were measured as were the extend of the protected areas. These figures served to determine the percentage of the remaining elephant range protected within each of the units.

Douglas-Hamilton, I., Burrill, A.
Using elephant carcass ratios to determine population trends. (1991)
African Wildlife: Research and Management, pp. 98-105

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Many African elephant populations have declined over the last two decades (Douglas-Hamilton. 1987). In most census zones, as the number of elephants decreases, the number of dead elephants increases. By counting both live and dead elephants a carcass ratio can be derived. This is the proportion of dead elephants to all elephants dead and live, and has been used as an index of relative elephant mortality (Douglas-Hamiton and Hillman, 1980).??In this paper successive counts of dead and live elephants in central, east, and southern Africa have been used to plot elephant trends against carcass ratios. It was found that the carcass ratio was correlated with the rate of decrease over a 4-yr period. The model was applied to new census data from Selous and Kilombero and found to give a close prediction of actual trends.

Douglas-Hamilton, I., Hillman, A.K.K.
ELEPHANT CARCASSES AND SKELETONS AS INDICATORS OF POPULATION TRENDS. (1981)
World Wildlife Fund /International Union for the Conservation of Nature Elephant Survey and Conservation Programme.

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Elephants have always posed problems to authorities, either as crop raiders, major agents of habitat change or as a potentially valuable natural resource. Decision-makers have had to consider such diverse questions as whether elephants should be culled as an ecological management policy within a park, or whether anti-poaching forces should be reinforced and the ivory trade banned to prevent their elimination. The true status of elephant populations has been confused by their apparent over-abundance in some areas and their disappearance from others. In every case where decisions must be made, knowledge is required of how many elephants there are and how their numbers are changing.

Douglas-Hamilton, I., Wittemyer, G., and Ihwagi, F. on behalf of Save the Elephants.
LEVELS OF ILLEGAL KILLING OF ELEPHANTS IN THE LAIKIPIA-SAMBURU MIKE SITE. (2010)
CITES CoP15 Inf. 40 (Rev. 1)

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A programme to monitor elephant mortality was agreed by the CITES parties in 1997. The MIKE (Monitoring of the Illegal Killing of Elephants) programme, was approved by CITES in 2000; and its current modus operandi follow Res. 10.10 (Rev. CoP14). It was set up to detect changes in levels of illegal killing of elephants. Kenya has been monitoring elephant mortality through the Kenya Wildlife Service of all elephant populations in the country since 1990, including the current Laikipia-Samburu MIKE site. The information is stored in a national elephant mortality database. A report summarizing all information from 1990 to 2002 was undertaken prior to the inception of MIKE, intended by Kenya as their own base line against which future change could be measured (Thouless et al. 2008).

Durant et al
Developing fencing policies for dryland ecosystems. (2015)
Journal of Applied Ecology 2015 doi: 10.1111/1365-2664.12415

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1. In dryland ecosystems, mobility is essential for both wildlife and people to access unpre-dictable and spatially heterogeneous resources, particularly in the face of climate change. Fences can prevent connectivity vital for this mobility. 2. There are recent calls for large-scale barrier fencing interventions to address human–wild-life conflict and illegal resource extraction. Fencing has costs and benefits to people and wild-life. However, the evidence available for facilitating sound decision-making for fencing initiatives is limited, particularly for drylands. 3. We identify six research areas that are key to informing evaluations of fencing initiatives: economics, edge permeability, reserve design, connectivity, ecosystem services and communities. 4. Policy implications.

Ganswindt, A., Muenscher, S., Henley, M., Henley, S., Heistermann, M., Palme, R., Thompson, P., Bertschinger, H.
Endocrine correlates of musth and the impact of ecological and social factors in free-ranging African elephants (Loxodonta africana) (2010)
Hormones and Behavior, Volume 57, Issues 4-5, April 2010, Pages 506-514. doi:10.1016/j.yhbeh.2010.02.009

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Sexual activity in mature male African elephants is predominantly associated with the occurrence of musth, a state or condition which refers to a set of physical, physiological and behavioral characteristics, including an elevation in androgen levels. Although musth appears to be energetically costly, the degree to which it is associated with changes in adrenal endocrine function (e.g., glucocorticoid output) is still unclear. To investigate the possible effect of musth on adrenocortical function, and the impact of socioecological changes on androgen and glucocorticoid levels, six adult African elephant bulls were followed for 13 months in the Kruger National Park, South Africa, and observations and fecal sample collection for endocrine monitoring was carried out about twice weekly.

Ganswindt, A., Münscher, S., Henley, M., Palme, R., Thompson, P., Bertschinger, H.
Concentrations of faecal glucocorticoid metabolites in physically injured free-ranging African elephants Loxodonta africana. (2010)
Wildlife Biology 16(3):323-332. doi: 10.2981/09-081

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Free-ranging African elephants Loxodonta africana use their front feet frequently during the process of foraging and this could be the reason for the high prevalence of physical injuries to these parts of the body. Although the occurrence of severe lameness caused by foot lesions in adult elephants has already been investigated and the clinical and pathological findings have been reported, the effect of foot injuries on glucocorticoid levels as a potential physiological stress response has not been examined. Given the practical difficulties involved in monitoring unpredictable events in free-ranging animals, like the occurrence of foot injuries in elephants, it is not surprising that information regarding the endocrine correlates of physical injury is still limited for elephants.

Ganswindt, A., Rasmussen, H., Heistermanna, M., Hodges, K.
The sexually active states of free-ranging male African elephants. (2005)
Hormones and Behavior 47 (2005) 83– 91, Elsevier Inc.

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Musth in male African elephants, Loxodonta africana, is associated with increased aggressive behavior, continuous discharge of urine, copious secretions from the swollen temporal glands, and elevated androgen levels. During musth, bulls actively seek out and are preferred estrous females although sexual activity is not restricted to the musth condition. The present study combines recently established methods fecal hormone analysis with long-term observations on male-female associations as well as the presence and intensity of physical signals to provide a more detailed picture about the physical, physiological, and behavioral characteristics of different states of sexual activity in free ranging African elephants.

Georgiadis, N.J., Ihwagi, F., Olwero, J.G.N., Roman, S.R.
Savanna herbivore dynamics in a livestock-dominated landscape. (2007)
Elsevier

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Conserving African wildlife in human-occupied landscapes requires management intervention that is guided by a mechanistic understanding of how anthropogenic factors influence large-scale ecological processes. In Laikipia District, a dry savanna region in northern Kenya where wildlife share the landscape with humans and livestock, we examined why five of nine wild ungulate species suffered protracted declines on properties receiving the greatest conservation investment. Of 10 alternative causes examined, only an increase in predation, interacting with brief periods of high and low rainfall, was consistent with the timing, synchrony, duration and species composition of observed ungulate declines.

Gessner, J., Buchwald, R., Wittemyer, G.
Assessing species occurrence and species-specific use patterns of bais (forest clearings) in Central Africa with camera traps. (2014)
African Journal of Ecology 52:59-68. DOI: 10.1111/aje.12084

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The impacts of increasing resource extraction on biodiversity in the Central African rainforest are largely unknown, in part due to the lack of baseline data on species occurrence across the basin. Natural forest clearings (bais) in this region are key habitats for a variety of vertebrates and offer opportunities for monitoring species distribution. Information on species composition, however, is lacking from the majority of areas (except for long-term study sites). Approaches and protocols for short-term bai assessments can greatly advance such baseline knowledge. This study demonstrates that camera traps provide an effective method for species inventories (species occurrence and temporal activity patterns) and monitoring at bais across the broader region. In comparison with direct observational studies, they performed especially well regarding rare and nocturnal species.

Goldenberg, S. Z., Douglas-Hamilton, I., Daballen, D. and Wittemyer, G.
Challenges of using behavior to monitor anthropogenic impacts on wildlife: a case study on illegal killing of African elephants (2017)
Anim Conserv, 20: 215–224. doi:10.1111/acv.12309

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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.

Goldenberg, S. Z., Douglas-Hamilton, I., Wittemyer, G.
Intergenerational change in African elephant range use is associated with poaching risk, primary productivity and adult mortality. (2018)
Proc. R. Soc. B 285: 20180286. http://dx.doi.org/10.1098/rspb.2018.0286

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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.

Goldenberg, S. Z., WITTEMYER, G.
Orphaned female elephant social bonds reflect lack of access to mature adults (2017)
SCIENTIFIC Reports | 7: 14408 | DOI:10.1038/s41598-017-14712-2

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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.

Goldenberg, S. Z., Wittemyer, G.
Orphaning and natal group dispersal are associated with social costs in female elephants (2018)
Elsevier Animal Behaviour Volume 143, September 2018, Pages 1-8 https://doi.org/10.1016/j.anbehav.2018.07.002

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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.

Goldenberg, S.Z, Oduor, S., Kinnaird, M.F., Daballen, D., Douglas‐Hamilton, I., Wittemyer, G.
Evidence of strong spatial segregation between elephant subpopulations in the contiguous Laikipia–Samburu ecosystem in Kenya (2016)
African Journal of Ecology, 54(2), 261-264 https://doi.org/10.1111/aje.12310

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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)

Goldenberg, S.Z., de Silva, S., Rasmussen, H.B., Douglas-Hamilton, I., and Wittemyer, G
Controlling for behavioural state reveals social dynamics among male African elephants, Loxodonta africana. (2014)
Animal Behaviour 95 (2014) 111-119

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The drivers of social affiliation may vary over time as individuals change their goals with respect to changing environments or physical condition. Studies of companion preference rarely consider shifts in motivational state, despite the potential importance of such shifts in structuring association and population processes. Ignoring state dependence in social behaviour may weaken the ability to recognize social properties and identify their underlying drivers. Modifying established approaches, we apply a state-specific analysis to investigate social properties in male African elephants, which are thought to be weakly social.

Goldenberg, S.Z., Douglas-Hamilton, I., Wittemyer, G
Vertical Transmission of Social Roles Drives Resilience to Poaching in Elephant Networks (2015)
Current Biology 26 (2016) pp. 75-79. Elsevier. DOI 10.1016/j.cub.2015.11.005

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Network resilience to perturbation is fundamental to functionality in systems ranging from synthetic communication networks to evolved social organization. While theoretical work offers insight into causes of network robustness, examination of natural networks can identify evolved mechanisms of resilience and how they are related to the selective pressures driving structure. Female African elephants (Loxodonta africana ) exhibit complex social networks with node heterogeneity in which older individuals serve as connectivity hubs. Recent ivory poaching targeting older elephants in a well studied population has mirrored the targeted removal of highly connected nodes in the theoretical literature that leads to structural collapse. Here we tested the response of this natural network to selective knockouts.

Goldenberg, S.Z., Hahn, N., Stacy-Dawes, J., Chege, S.M., Daballen, D., Douglas-Hamilton, I., Lendira, R.R., Lengees, M.J., Loidialo, L.S., Omengo, F. and Pope, F.
Movement of rehabilitated African elephant calves following soft release into a wildlife sanctuary (2022)
Frontiers in Conservation Science, p.55.

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The ability to locate essential resources is a critical step for wildlife translocated into novel environments. Understanding this process of exploration is highly desirable for management that seeks to resettle wildlife, particularly as translocation projects tend to be expensive and have a high potential for failure. African savannah elephants (Loxodonta africana) are very mobile and rely on large areas especially in arid environments, and are translocated for differing management and conservation objectives. Thus, research into how translocated elephants use the landscape when released may both guide elephant managers and be useful for translocations of other species that adjust their movement to social and ecological conditions. In this study, we investigated the movement of eight GPS tracked calves (translocated in three cohorts) following their soft release into a 107 km2 fenced wildlife sanctuary in northern Kenya and compared their movement with that of five tracked wild elephants in the sanctuary. We describe their exploration of the sanctuary, discovery of water points, and activity budgets during the first seven, 14, and 20 months after release.

Graham, M.
Coexistence in a land use mosaic? Land use, risk and elephant ecology in Laikipia District, Kenya. (2006)
University of Cambridge for the degree of Doctor of Philosophy

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This thesis is about the patterns, determinants and consequences of human-elephant interaction in Laikipia District in northern Kenya. Laikipia is located outside of formally protected areas, supports a range of land use types and harbours Kenya’s second largest elephant population comprised of over 3,000 animals. I use interdisciplinary methods and multiple scales of spatial analysis to examine elephant distribution, persistence and interactions with people in this human landscape. At a course scale, results from several data sources show that elephants occur across almost 50% of Laikipia District and, intriguingly, are relatively evenly distributed across locations under cultivation, settlement and livestock production.

Graham, M.D., Douglas-Hamilton, I., Adams, W.M., Lee, P. C.
The movement of African elephants in a human-dominated land-use mosaic. (2009)
Animal Conservation 12 (2009) 445–455. doi:10.1111/j.1469-1795.2009.00272.x  

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Land outside of gazetted protected areas is increasingly seen as important to the ?future of elephant persistence in Africa. However, other than inferential studies on ?crop raiding, very little is understood about how elephants Loxodonta africana use ?and are affected by human-occupied landscapes. This is largely a result of ?restrictions in technology, which made detailed assessments of elephant movement? outside of protected areas challenging. Recent advances in radio telemetry have ?changed this, enabling researchers to establish over a 24-h period where tagged ?animals spend their time. We assessed the movement of 13 elephants outside of ?gazetted protected areas across a range of land-use types on the Laikipia plateau in? north-central Kenya.

Graham, M.D., Gichohi, N., Kamau, F. Aike, G., Craig, B., Douglas-Hamilton, I. and Adams, W.
The Use of Electrified Fences to Reduce Human Elephant Conflict: A Case Study of the Ol Pejeta Conservancy, Working Paper 1. (2009)
Laikipia Elephant Project, Nanyuki, Kenya

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Human-elephant conflict is a significant problem in Africa and Asia, particularly where land? managed for conservation adjoins land under cultivation. Electrified fences are increasingly used? to reduce such conflict by preventing access by elephants to vulnerable land. However, despite ?the growing number of electrified fences erected to address human-elephant conflict, there ?have been few empirical studies of their effectiveness. Here we assess the performance of an? electrified fence constructed around the 370km2 Ol Pejeta Conservancy on the Laikipia Plateau ?in north-central Kenya.

Greyling, M.
Sex and age-related distinctions in the feeding ecology of the African Elephant, Loxodonta africana. (2004)
PhD thesis University of theWitwatersrand, Johannesburg

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Sexual dimorphism in size in the African savanna elephant, Loxodonta africana, is pronounced. Allometric differences between the sexes lead to dissimilar nutritional demands, which result in sex related distinctions in feeding ecology. This extension of the Jarman-Bell Principle to an intra-specific level has been referred to as the Body Size Hypothesis (BSH).??This study established whether different nutritional requirements of elephant size/sex classes resulted in functional distinctions in feeding ecology between elephant bull groups and family units. Plant based surveys on woody species were conducted at the feeding sites of both bull groups and family units of elephants during the dry season period of resource limitation within the Associated Private Nature Reserves of South Africa.

Greyling, M.D., Ford, M., Potgieter, H. C., van Aarde, R.J.
Influence of Gestation on Uterine Endometrial Steroid Receptor Concentrations in the African Elephant, Loxodonta africana’. (1998)
Biology of Reproduction 58, 60-64

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The modulatory effects of gestational age and circulating concentrations of progesterone, 5-pregnane-3,20-dione, and estradiol-17 on the uterine sex steroid hormone receptor levels of the African elephant were investigated. Uterine tissue biopsies and blood samples were obtained from animals culled in the Kruger National Park. Estrogen and progesterone receptor concentrations were determined in uterine biopsies from subadult, lactating, early-, mid-, and late-pregnant elephants, by equilibrium binding assays. Circulating estradiol-171 and progesterone concentrations were measured by means of RIAs, while plasma concentrations of 5-pregnane-3,20-dione were determined with an amplified ELISA. Significant inverse correlations of the concentrations of estrogen and progesterone receptors with the gestational stage of the elephants were observed.

Greyling, M.D., McCay, M., Douglas-Hamilton, I.
Green Hunting as an Alternative to Lethal Hunting. (2004)
Save the Elephants

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Private nature reserves and adjoining large national parks such as the 2.2 million ha Kruger National Park (KNP) in South Africa, not only need to function as integrated ecological units but also need to be financially viable to support efficient management practices. The Association of Private Nature Reserves (APNR), on the western boundary of the KNP encompass an area of 180 000 ha and forms one of the largest private nature reserves in the world. These reserves host from time to time some of the few remaining large tusked bulls, which periodically emerge from the KNP. As hunting is permitted within the APNR, trophy bulls are of economic importance to the reserves. Green hunting of elephant bulls was pioneered by Save the Elephants within the Timbavati Private Nature Reserve in 1998.

Greyling, M.D., van Aarde, R.J., Potgieter, H. C.
Ligand specificity of uterine oestrogen and progesterone receptors in the subadult African elephant, Loxodonta africana (1997)
Journal of Reproduction and Fertility. 1997. 109. 199-204

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Relative high annual rates of population growth in African elephants confined to conservation areas in southern Africa (for example, 7%; Hall-Martin, 1992) may have negative consequences for the maintenance of biological diversity. As a result the densities of some of these populations are reduced artificially through culling operations that are opposed both by some conservationists and the concerned public. There is, therefore, a need for the development of alternative and generally acceptable techniques for inhibiting population growth. In this regard Poole (1993) and Short (1992) suggested that consideration should be given to the use of contraceptives or contragestins, such as an antiprogestin, like mifepristone (RU?486; Roussel Uclaf, Paris), that may block uterine receptor activity. However, the use of such treatments require detailed knowledge of the reproductive endocrinology in the elephant.

Hahn, N. R., Wall, J., Denninger-Snyder, K., Goss, M., Sairowua, W., Mbise, N., Estes, A. B., Ndambuki, S., Mjingo, E. E., Douglas-Hamiliton, I., Wittemyer, G.
Risk perception and tolerance shape variation in agricultural use for a transboundary elephant population (2022)
Journal of Animal Ecology, 91, 112– 123. https://doi.org/10.1111/1365-2656.13605

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Abstract To conserve wide-ranging species in human-modified landscapes, it is essential to understand how animals selectively use or avoid cultivated areas. Use of agriculture leads to human–wildlife conflict, but evidence suggests that individuals may differ in their tendency to be involved in conflict. This is particularly relevant to wild elephant populations. We analysed GPS data of 66 free-ranging elephants in the Serengeti-Mara ecosystem to quantify their use of agriculture. We then examined factors influencing the level of agricultural use, individual change in use across years and differences in activity budgets associated with use. Using clustering methods, our data grouped into four agricultural use tactics: rare (12.8%; 9%). Sporadic and seasonal individuals represented two-thirds (67%) of recorded GPS fixes in agriculture, compared to 32% from habitual individuals. Increased agricultural use was associated with higher daily distance travelled and larger home range size, but not with age or sex. Individual tactic change was prevalent and the habitual tactic was maintained in consecutive years by only five elephants. Across tactics, individuals switched from diurnal to nocturnal activity during agricultural use, interpreted as representing similar risk perception of cultivated areas. Conversely, tactic choice appeared to be associated with differences in risk tolerance between individuals. Together, our results suggest that elephants are balancing the costs and benefits of crop usage at both fine (e.g. crop raid events) and long (e.g. yearly tactic change) temporal scales. The high proportion of sporadic and seasonal tactics also highlights the importance of mitigation strategies that address conflict arising from many animals, rather than targeted management of habitual crop raiders. Our approach can be applied to other species and systems to characterize individual variation in human resource use and inform mitigations for human–wildlife coexistence.

Hahn, N.R., Bombaci, S.P. & Wittemyer, G.
Identifying conservation technology needs, barriers, and opportunities (2022)
Sci Rep 12, 4802 (2022). https://doi.org/10.1038/s41598-022-08330-w

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Amid accelerating threats to species and ecosystems, technology advancements to monitor, protect, and conserve biodiversity have taken on increased importance. While most innovations stem from adaptation of of-the-shelf devices, these tools can fail to meet the specialized needs of conservation and research or lack the support to scale beyond a single site. Despite calls from the conservation community for its importance, a shift to bottom-up innovation driven by conservation professionals remains limited. We surveyed practitioners, academic researchers, and technologists to understand the factors contributing to or inhibiting engagement in the collaborative process of technology development and adoption for feld use and identify emerging technology needs. High cost was the main barrier to technology use across occupations, while development of new technologies faced barriers of cost and partner communication. Automated processing of data streams was the largest emerging need, and respondents focused mainly on applications for individual-level monitoring and automated image processing. Cross-discipline collaborations and expanded funding networks that encourage cyclical development and continued technical support are needed to address current limitations and meet the growing need for conservation technologies.

Hahn, N.R., Wall, J., Denninger‐Snyder, K., Goss, M., Sairowua, W., Mbise, N., Estes, A.B., Ndambuki, S., Mjingo, E.E., Douglas‐Hamiliton, I. and Wittemyer, G.
Risk perception and tolerance shape variation in agricultural use for a transboundary elephant population. (2022)
Journal of Animal Ecology, 91(1), pp.112-123.

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Abstract To conserve wide-ranging species in human-modified landscapes, it is essential to understand how animals selectively use or avoid cultivated areas. Use of agriculture leads to human–wildlife conflict, but evidence suggests that individuals may differ in their tendency to be involved in conflict. This is particularly relevant to wild elephant populations. We analysed GPS data of 66 free-ranging elephants in the Serengeti-Mara ecosystem to quantify their use of agriculture. We then examined factors influencing the level of agricultural use, individual change in use across years and differences in activity budgets associated with use. Using clustering methods, our data grouped into four agricultural use tactics: rare (12.8%; 9%). Sporadic and seasonal individuals represented two-thirds (67%) of recorded GPS fixes in agriculture, compared to 32% from habitual individuals. Increased agricultural use was associated with higher daily distance travelled and larger home range size, but not with age or sex. Individual tactic change was prevalent and the habitual tactic was maintained in consecutive years by only five elephants. Across tactics, individuals switched from diurnal to nocturnal activity during agricultural use, interpreted as representing similar risk perception of cultivated areas. Conversely, tactic choice appeared to be associated with differences in risk tolerance between individuals. Together, our results suggest that elephants are balancing the costs and benefits of crop usage at both fine (e.g. crop raid events) and long (e.g. yearly tactic change) temporal scales. The high proportion of sporadic and seasonal tactics also highlights the importance of mitigation strategies that address conflict arising from many animals, rather than targeted management of habitual crop raiders. Our approach can be applied to other species and systems to characterize individual variation in human resource use and inform mitigations for human–wildlife coexistence.

Hart, J., Gobush, K., Maisels, F., Wasser, S., Okita-Ouma, B., & Slotow, R.
African forest and savannah elephants treated as separate species (2021)
Oryx , Volume 55 , Issue 2 , March 2021 , pp. 170 - 171 DOI: https://doi.org/10.1017/S0030605320001386

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The African Elephant Specialist Group (AfESG) of IUCN will now treat African elephants as two species: the forest elephant Loxodonta cyclotis and the savannah elephant Loxodonta africana. This will be reflected in IUCN's Red List assessment update for African elephants, and in the next iteration of the African Elephant Status Report, both to be published in 2021. This concurs with Wilson & Reader (Mammal Species of the World, 2005), the primary IUCN reference on mammalian taxonomy, Wittemyer (in Handbook of the Mammals of the World, 2011), and Tassy & Shoshani (in Mammals of Africa, 2013).

Ihwagi et al.
Using Poaching Levels and Elephant Distribution to Assess the Conservation Efficacy of Private, Communal and Government Land in Northern Kenya. (2015)
PLoS ONE 10(9): e0139079. doi:10.1371/journal.pone.0139079

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Efforts to curb elephant poaching have focused on reducing demand, confiscating ivory and boosting security patrols in elephant range. Where land is under multiple uses and ownership, determining the local poaching dynamics is important for identifying successful conservation models. Using 2,403 verified elephant, Loxodonta africana, mortality records collected from 2002 to 2012 and the results of aerial total counts of elephants conducted in 2002, 2008 and 2012 for the Laikipia-Samburu ecosystem of northern Kenya, we sought to determine the influence of land ownership and use on diurnal elephant distribution and on poaching levels.We show that the annual proportions of illegally killed (i.e., poached) elephants increased over the 11 years of the study, peaking at 70% of all recorded deaths in 2012.

Ihwagi, F.I., Skidmore, A.K., Wang, T., Bastille‐Rousseau, G., Toxopeus, A.G., Douglas‐Hamilton, I.
Poaching Lowers Elephant Path Tortuosity: Implications for Conservation (2019)
The Journal of Wildlife Management 1–10; 2019; DOI: 10.1002/jwmg.21688

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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.

Ihwagi, F.W
LIVING IN A RISKY LANDSCAPE: ELEPHANT MOVEMENT IN RESPONSE TO POACHING (2018)
PhD Thesis. University of Twente

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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.

Ihwagi, F.W.
Forage Quality And Bark Utilisation By The African Elephant (Loxodonta africana) In Samburu And Buffalo Springs National Reserves, Kenya (2007)
M.Sc. Thesis, University of Nairobi

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Foraging behavior of elephants with respect to debarking of woody species was?investigated in Samburu and Buffalo Springs National Reserves, Kenya. Acacia elatior was?the most preferred species followed by Acacia tortilis. Both A. elatior and A. tortilis? dominate the woody vegetation accounting for over 80% of all woody plants. Debarking ?levels varied in different parts of the reserves and this was attributed to elephants’ densities ?and the ultimate influence of endaphic factors on species assemblages. Species diversity ?indices were negatively correlated with salinity indicating a direct influence of salinity on ?plant community structure. Both Acacia tortilis and A. elatior have the highest tolerance to? salinity and occur almost exclusively in saline areas.

Ihwagi, F.W., Chira, R.M., Kironchi, G., Vollrath, F. and Douglas-Hamilton, I.
Rainfall pattern and nutrient content influences on African elephants’ debarking behaviour in Samburu and Buffalo Springs National Reserves, Kenya (2021)
African Journal of Ecology, 50, 152-159. https://onlinelibrary.wiley.com/doi/abs/10.1111/j.1365-2028.2011.01305.x

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Abstract The magnitude of debarking by elephants was investigated in Samburu and Buffalo Springs National Reserves. About 1617 plants were monitored for debarking intensities for 6months spanning through dry and wet seasons. Debarking indices ranged from no debarking at all during the wet months to complete stem girding at the height of the dry season. A negative correlation was found between rainfall and debarking indices. It was hypothesized that nutrient content of the bark influenced the magnitude to which trees were debarked. Bark samples were collected from least, moderate and intensely debarked plants throughout the 6 months. These were analysed for cal- cium (Ca), sodium (Na), phosphorus (P), magnesium (Mg), potassium (K), nitrogen (N), iron (Fe), copper (Cu), man- ganese (Mn) and zinc (Zn). Significant positive correlations were found between debarking intensity and each of the nutrients N [crude protein (CP)], P, K and Zn. Bark was found to be richest in CP and Calcium. Neutral detergent fibre content was on average 67%. Monthly variations in nutrient composition were minimal. Acacia elatior, the most preferred species had significantly higher quantities of each of the four elements N, P, K and Zn than Acacia tortilis, the second most preferred woody species.

Ihwagi, F.W., Douglas-Hamilton, I.
A Report on the Samburu October 2005 Aerial Total Count. (2005)
Save the Elephants

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The count was conducted under the auspices of Northern Rangelands Trust (NRT) jointly funded by NRT, African Wildlife Foundation, Disney Foundation and Save the Elephants. Participants were drawn from the organization and a full list of individuals who took part is in the acknowledgement section. In particular, Onesmas Kahindi took a leading role in training and coordinating the census crew while and Rose Mayienda put the data together and made a single shape file of it. The report describes the methodology used and summarizes the results of the counts in form of tables and Maps.

Ihwagi, F.W., Thouless, C, Wang, T., Skidmore, A.K., Omondi, P., Douglas-Hamilton, I.
Night-day speed ratio of elephants as indicator of poaching levels (2017)
Ecological Indicators. Volume 84, January 2018, Pages 38–44 https://doi.org/10.1016/j.ecolind.2017.08.039

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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.

Ihwagi, F.W., Vollrath, F., Chira, R.M., Douglas-Hamilton, I., Kironchi, G.
The impact of elephants, Loxodonta africana, on woody vegetation through selective debarking in Samburu and Buffalo Springs National Reserves, Kenya (2009)
Blackwell Publishing Ltd, Afr. J. Ecol. doi: 10.1111/j.1365-2028.2009.01089.x

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The impact of elephants on the woody plant community through debarking was investigated in Samburu and Buffalo Springs National Reserves, Kenya. Acacia elatior Brenan, the most abundant tree species inthe riverine zone, accounted for 68%(n = 1375) of woody plants. A. tortilis (Forsskal) Hayne dominated plots away from the river. Debarking incidences were significantly higher for A. elatior than for other species indicating selective utilization. The riverine zone by virtue of having more trees of the preferred species, A. elatior, had the highest debarking incidences. Presence of very few saplings along the river is attributed to both elephant trampling and herb ivory by other species. An estimated 38.5% and 22.5% of the riverine A. elatior and A. tortilis trees respectively, were bound to die within the next 4–5 years because of severe debarking, ‡75% of bark circumference.

Johnson, A. S.
The Effects of Tactile and Visual Deterrents on Honey Badger Predation of Beehives (2018)
MSc Thesis Submitted in partial fulfillment of the requirements for the degree of Master of Arts Animal Behavior and Conservation, Hunter College The City University of New York

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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.

Kahindi, O.
Cultural perceptions of elephants by the Samburu people in northern Kenya (2001)
M.Sc. Thesis. Strathclyde University, Scotland

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The Samburu people of northern Kenya have co-existed with elephants since time immemorial.  The Samburu-elephant co-existence is facilitated by local knowledge gained through real experiences from direct interactions with, and actual observation of the elephant’s natural behaviour. The experiences are interpreted and coded through existing traditional belief systems and permeated to the community and descending generations through the vibrant oral system in the society.  The knowledge is an integral part of the co-existence. The Samburu perceive elephants in terms of individuals and individual groups rather than a population.  Individual elephants have meaningful and significant characters.  The Samburu perception is different from that of other organizations, past and present, interested in the elephants inhabiting Samburu District.

Kahindi, O., Wittemyer, G., King, J., Ihwagi, F., Omondi, P., Douglas-Hamilton, I.
Employing participatory l killing of elephants across diverse land uses in Laikipia–Samburu, Kenya (2009)
Blackwell Publishing Ltd, Afr. J. Ecol. doi: 10.1111/j.1365-2028.2009.01200.x

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Levels and trends of illegal killing of elephants are measured by the Convention on International Trade in Endangered Species (CITES) Monitoring the Illegal Killing of Elephants (MIKE) programme in sites across Africa and Asia. In the mostly unprotected Laikipia–Samburu MIKE site in northern Kenya, elephant mortality data were collected using both standard law enforcement monitoring procedures, relying on patrolling, and participatory methods involving local communities. Qualitatively, traditional patrolling techniques were more successful in protected areas whereas participatory approaches provided more information outside protected areas, where elephant were most at risk from ivory poachers. A minimum of 35% of the 389 verified carcasses during 2001–2003 were illegally killed.

Kahumbu, P. G., Douglas-Hamilton, I.
Elephant movements, home ranges and diurnal activity. (2002)
Chapter in A Dissertation presented to the Faculty of Princeton University

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As human encroachment into elephant rangelands restricts elephant movements, it is increasingly critical to identify elephant range needs, and the effects of barriers to movements for the purpose of planning conservation areas. Home ranges and movements were documented for six African elephants in a small protected area comprising mixed tropical forest, woodland and open habitats. Data were collected using four conventional VHF radio tracking system (Telonics) collars on two cows and two bulls, and LOTEK GPS radio collars on one bull and one cow. None of the collars performed as well as was expected, however, the LOTEK GPS collars provided far superior data to the conventional VHF system.

Kantai, R
Tracing Community Conservation Evolution, Power Relations and Sustainability in Northern Kenya. (2012)
MSc. Thesis, University of Oxford

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Abstract The narrative of Fortress Conservation was based on strategies dominated by attempts to reserve places for nature, to separate humans from nature, and to prevent consumptive use or other forms of human impact. The counter-narrative of Community Conservation promises to reconcile conservation and development objectives, and ensure the interests of the local people are taken into account. This narrative is no panacea however, and in the evolution of community conservation in Kenya, several initiatives have fallen victim to poor governance or the lack thereof, or worse, the retrogression into the colonial model of conservation through exclusion. A new model of community conservation in Kenya is emerging, distinguished by the governance and leadership of umbrella bodies and land owner associations, state involvement and community buy-in in a new frontier of conservation on communal lands.

King L.E., Lawrence, A., Douglas-Hamilton, I., Vollrath, F.
Beehive fence deters crop-raiding elephants. (2009)
Blackwell Publishing Ltd, Afr. J. Ecol., 47, 131–137

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Previous work has shown that African elephants Loxodonta africana will avoid African honeybees Apis mellifera scutellata. Here we present results from a pilot study conducted to evaluate the concept of using beehives to mitigate elephant crop depredation. In Laikipia, Kenya, we deployed a 90-m fence-line of nine inter-connected hives, all empty, on two exposed sides of a square two-acre farm that was experiencing high levels of elephant crop depredation. Compared with a nearby control farm of similar status and size, our experimental farm experienced fewer raids and consequently had higher productivity.??Socioeconomic indicators suggest that not only was the concept of a beehive fence popular and desired by the community but also that it can pay for its construction costs through the sale of honey and bee products.

King, L., Pardo, M., Weerathunga, S., Kumara, T.V., Jayasena, N., Soltis, J. and de Silva, S
Wild Sri Lankan elephants retreat from the sound of disturbed Asian honey bees. (2018)
Current Biology 28, R51–R65, January 22. DOI: 10.1016/j.cub.2017.12.018 http://www.cell.com/current-biology/abstract/S0960-9822(17)31649-4

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King, L., Raja, N., Kumar, M., Heath, N. and Pope, F.
Part I: Development of a new Human-Elephant Coexistence Toolbox for communities living with African savannah elephants (Loxodonta Africana) (2022)
Pachyderm, 63, pp.153-157.

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Introduction: This field note is to invite our colleagues to peer review and test a new illustrated Human-Elephant Coexistence (HEC) Toolbox that is being developed in Kenya by Save the Elephants (STE) under the organizations’ mission to secure a future for elephants and to develop a tolerant relationship between humans and elephants. Through presenting the first edition here (Fig. 1), we are inviting our elephant colleagues and community leaders from across the African savannah elephant range States to provide feedback, or any corrections, on the tools, as well as sharing content for additional methods not yet represented. By publishing our process and methods for how we are compiling this encyclopaedia of HEC tools and this novel approach to the peer review process, we hope to provide a transparent process to gauging the validity of the methods presented. This is particularly important because some of the technical advice around the conflict reduction tools presented are not published formally in the scientific literature

King, L.E.
Could honey bees be effective deterrents for Asia’s crop-raiding elephants? (2013)
Sanctuary Asia, October 2013

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As our ancient jeep bucked and jolted to a stop on the steep bank, I caught a glimpse of the thickly forested edge of Wasgamuwa National Park and wondered, not for the fi rst time on this trip, if some of the wild elephants that inhabited this part of the island were surreptitiously watching our arrival. We waited silently for a flicker of elephant movement. Compared with my Samburu study site in Kenya, where elephants are plentiful and habituated to my vehicle, I was unused to the need to exercise such patience, so I enjoyed the time mulling over all that I had learnt about Sri Lankaâ??s elephants.

King, L.E.
Bees help Protect Crops From Damage by Elephants in Kenya. (2009)
Bee Craft. September 2009. P9-11

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The Elephants and Bees Research project is one of Save the Elephants' innovative programmes designed to explore the natural world for solutions to human-elephant conflict. The project uses in-depth knowledge and observation of elephant behaviour to reduce damage from crop-raiding elephants, using African honey bees. Save the Elephants is a research team headed by the elephant expert Dr. Iain Douglas-Hamilton, OBE. In collaboration with the Kenya Wildlife Service, we are investigating crop protection methods that can be financed and managed by the farmers themselves to provide long term solutions to human-elephant conflict.

King, L.E.
The interaction between the African elephant (Loxodonta africana africana) and the African honey bee (Apis mellifera scutellata) and its potential application as an elephant deterrent. (2010)
A thesis submitted to the University of Oxford for the degree of Doctor of Philosophy in Zoology, Balliol College.

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The conceptual origin of this DPhil thesis was based on one foundation publication by Vollrath and Douglas-Hamilton (2002a) “African bees to control African elephants”. The authors made a unique discovery that African elephants will avoid feeding on acacia trees that host beehives, either empty or occupied by African honey bees. The concept that elephants might hold a long term memory about bees that could be so negative as to evolve avoidance behaviour towards an otherwise favourite food source, was deeply intriguing and warranted further study. Two key research questions are asked in this thesis and are reflected in the title (i) what happens when elephants and honey bees interact and (ii) how can we adapt this behaviour into a potential deterrent system for crop-raiding elephants. Hence, this is really a thesis of two halves blending both disciplines of natural and social sciences.

King, L.E., Douglas-Hamilton, I., Vollrath, F.
Beehive fences as effective deterrents for crop-raiding elephants: field trials in nothern Kenya (2011)
African Journal of Ecology. doi: 10.1111/j.1 365-2028.2011.01275.x

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Increasing elephant populations in Kenya since 1989 have been widely praised as a conservation success story. However, where elephants and agricultural land overlap, incidents of human-elephant conflict are on the increase. Wildlife managers and farmers are now trying different farm-based deterrents to keep elephants out of crops. Here, we present data on the effectiveness of a novel beehive fence deployed in a Turkana community of 62 communally run farms in Kenya. Specifically, 1700m of beehive fences semi-sorrounded the outer boundaries of seventeen farms, and we compared elephant farm invasion events with these and to seventeen neighboring farms whose boundaries were 'protected' only by thorn bush barriers. We present data from 45 farm invasions, or attempted invasions, recorded over 2 years. Thirteen groups of elephants approached beehive fences and turned away.

King, L.E., Douglas-Hamilton, I., Vollrath, F.
African elephants run from the sound of disturbed bees. (2007)
Current Biology Vol 17 No 19. R832

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Encroaching human development into former wildlife areas [1] is compressing African elephants into ever smaller home ranges, causing increased levels of human– elephant conflict [2]. African honeybees have been proposed as a possible deterrent to elephants [3]. We have performed a sound playback experiment to study this hypothesis. We found that a significant majority of elephants, in a sample of 18 well-known families and subgroups of varying sizes, reacted negatively — immediately walking or running away — when they heard the buzz of disturbed bees, while they ignored the control sound of natural white-noise. Whether the observed response was the result of individual conditioning or of learning by social facilitation remains to be established. Our study strongly supports the hypothesis that bees — and perhaps even their buzz alone — may be deployed to keep elephants at bay.

King, L.E., Lala, F., Nzumu, H., Mwambingu, E., and Iain Douglas-Hamilton, I.
Beehive fences as a multidimensional conflict-mitigation tool for farmers coexisting with elephants (2017)
Conservation Biology, Volume 00, No. 0, 1–10 DOI: 10.1111/cobi.12898

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Abstract: Increasing habitat fragmentation and human population growth in Africa has resulted in an escalation in human–elephant conflict between small-scale farmers and free-ranging African elephants (Loxodonta Africana). In 2012 Kenya Wildlife Service (KWS) implemented the national 10-year Conservation and Management Strategy for the Elephant in Kenya, which includes an action aimed at testing whether beehive fences can be used to mitigate human–elephant conflict. From 2012 to 2015, we field-tested the efficacy of beehive fences to protect 10 0.4-ha farms next to Tsavo East National Park from elephants. We hung a series of beehives every 10 m around the boundary of each farm plot. The hives were linked with strong wire.

King, L.E., Soltis, J., Douglas-Hamilton, I., Savage, A., Vollrath, F.
Bee Threat Elicits Alarm Call in African Elephants. (2010)
PLoS ONE 1 April 2010 | Volume 5 | Issue 4 | e10346

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Unlike the smaller and more vulnerable mammals, African elephants have relatively few predators that threaten their?survival. The sound of disturbed African honeybees Apis meliffera scutellata causes African elephants Loxodonta africana to?retreat and produce warning vocalizations that lead other elephants to join the flight. In our first experiment, audio?playbacks of bee sounds induced elephants to retreat and elicited more head-shaking and dusting, reactive behaviors that?may prevent bee stings, compared to white noise control playbacks. Most importantly, elephants produced distinctive?‘‘rumble’’ vocalizations in response to bee sounds. These rumbles exhibited an upward shift in the second formant location,?which implies active vocal tract modulation, compared to rumbles made in response to white noise playbacks.

Kinyanjui, M.W., Raja, N.R.,  Ewan J Brennan, E. J., King, L. E., Tiller, L. N
Local attitudes and perceived threats of human-elephant conflict: a case study at Lake Jipe, Kenya (2020)
Pachyderm No. 61 July 2019–June 2020

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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.

Koskei M., Okita-Ouma B., Lala F., Mwazo A., Kibara D., Tiller L., King L., Pope F. & Douglas-Hamilton I.
The effect of the new standard gauge railway (SGR) on elephant movement in Tsavo Ecosystem, Kenya (March 2016 – March 2018) (2018)
Save The Elephants & Kenya Wildlife Service

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In 2014, the Government of Kenya initiated construction of the Standard Gauge Railway (SGR) linking Kenya’s largest port city, Mombasa, and her capital city, Nairobi. The construction of the Mombasa – Nairobi SGR was completed and officially launched for use in June 2017. The SGR cuts through the 23,000 Km2 Tsavo National Parks comprising Tsavo East and Tsavo West National Parks, and is home to the largest single elephant population in Kenya, numbering approximately 13,000 animals (2017 total aerial count) as well as other mammalian species. Save the Elephants (STE) in partnership with Kenya Wildlife Service (KWS) fitted radio tracking collars on ten Tsavo elephants in March 2016. The main objective of this project was to understand the potential impact of the new SGR and highways in Tsavo on elephant movement and the effectiveness of the mitigation measures put in place to maintain ecosystem connectivity. A further 20 radio tracking collars deployed in February 2018 is expected to enrich these data we are collecting. However, this report documents findings from the initial ten collars deployed in March 2016. In this report, we explore movement patterns and hotspots of elephants’ use in relation to the new SGR and the Voi – Taveta highway after two years of tracking. The home ranges of some elephants crossing the SGR extends from Yatta plateau in Tsavo East to beyond Mkomazi National Park in Tanzania. Out of the eight elephants collared along the SGR; six of them (some with their families), have managed to cross the SGR during the two years of monitoring. The two most frequently used SGR crossing points for the tracked elephants in Tsavo were the Maungu corridor and at the Manyani corridor. The two corridors are one of six SGR underpasses designed specifically for wildlife use. The monitoring of wildlife utilization of SGR crossing underpasses by KWS patrols teams have shown that, through direct animal signs such as footprints and dung, elephants are learning relatively quicker to use the underpasses than other wildlife species. Movement patterns of the collared elephants also show that the erection of double electric fence, SGR fence and David Sheldrick Wildlife Trust’s fence at Ngutuni, has completely blocked elephant movements to the high-usage area between the SGR and the Mombasa highway. Elephants attempting to cross over the SGR and highway from Ngutuni Conservancy towards the Sagalla farms have also had their movements curtailed. The monitoring of the SGR, on foot or by vehicle, for wildlife crossing shows that elephant movements have been greatly impacted by the fragmentation of the habitat and blocking of normal migration routes. Some of the tracked elephants spent more than a year moving up and down along the SGR, but were not able to cross it after two years. The analysis of the mean speed in elephant tracking data indicates that elephants are slower crossing the existing Mombasa highway than the new raised SGR. One of the major challenge facing the underpass utilization by wildlife is the proliferation of illegal human settlements along the SGR and near the main underpasses. This applies especially to the Voi – Bachuma section of the SGR, around Ndara and Maungu areas. The other notable threat to wildlife use of the SGR underpasses is the high influx of livestock recorded going into the park through those underpasses. We also identified in this study that elephants are attracted to the borrow pits not completely filled after construction, along the SGR. The borrow pits collects and avail water to elephants and other wildlife species, but may expose them to potential train and vehicle accidents.

Koskei, M., Kolowski, J., Wittemyer, G., Lala, F., Douglas-Hamilton, I. and Okita-Ouma, B.
The role of environmental, structural and anthropogenic variables on underpass use by African savanna elephants (Loxodonta africana) in the Tsavo Conservation Area. (2022)
Global Ecology and Conservation, 38, p.e02199.

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A B S T R A C T: Wildlife crossing structures are effective interventions for mitigating fragmentation of habitats by linear infrastructure. The 2017 construction of a new railway cutting through the Tsavo Conservation Area (TCA), home to the largest elephant population in Kenya, affected wildlife movement and habitat connectivity. Although numerous studies have investigated the use of wildlife crossing structures by a wide range of species, few have focused on their use by megaherbivores. In this study, we examined use of 41 wildlife crossing structures by African savanna elephants (Loxodonta africana) along a 133 km section of new railway in Tsavo, Kenya. We used a generalized linear mixed modeling approach to assess the relationship between elephant crossing rate over 28 months between July 2017 to April 2021 and explanatory factors including crossing structure attributes, livestock presence and proximity to highways, water points and human settlement. We found that structural attributes of crossing structures were most strongly associated with the elephant crossing rate, particularly height and its interaction with type of crossing structure (bridges, wildlife underpasses and culverts). Higher crossing structures were associated with higher crossing rate, with the largest influence of height at culverts and wildlife underpasses. Although bridges comprised only 19.5 % of the 41 available crossing structures, they accounted for a disproportionately high number of elephants crossing events (56 %). The results demonstrated the importance of bridges over designated crossing structures for elephants, with predicted seasonal counts of elephant crossings being 0.31 for average sized culverts, 2.88 for wildlife underpasses and 5.86 for bridges. The environmental and anthropogenic variables were not strongly associated with elephant crossing rate. Our findings have direct application for future siting and design of crossing structures across elephant ranges.

Kuiper, T., Altwegg, R., Beale, C., Carroll, T., Dublin, H.T., Hauenstein, S., Kshatriya, M., Schwarz, C., Thouless, C.R., Royle, A. and Milner-Gulland, E.J.
Drivers and facilitators of the illegal killing of elephants across 64 African sites (2023)
Proceedings of the Royal Society B, 290(1990), p.20222270.

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Ivory poaching continues to threaten African elephants. We (1) used criminology theory and literature evidence to generate hypotheses about factors that may drive, facilitate or motivate poaching, (2) identified datasets representing these factors, and (3) tested those factors with strong hypotheses and sufficient data quality for empirical associations with poaching. We advance on previous analyses of correlates of elephant poaching by using additional poaching data and leveraging new datasets for previously untested explanatory variables. Using data on 10 286 illegally killed elephants detected at 64 sites in 30 African countries (2002–2020), we found strong evidence to support the hypotheses that the illegal killing of elephants is associated with poor national governance, low law enforcement capacity, low household wealth and health, and global elephant ivory prices. Forest elephant populations suffered higher rates of illegal killing than savannah elephants. We found only weak evidence that armed conflicts may increase the illegal killing of elephants, and no evidence for effects of site accessibility, vegetation density, elephant population density, precipitation or site area. Results suggest that addressing wider systemic challenges of human development, corruption and consumer demand would help reduce poaching, corroborating broader work highlighting these more ultimate drivers of the global illegal wildlife trade.

Kuriyan, R.
Linking local perceptions of elephants and conservation: Samburu pastoralists in Northern Kenya. (2002)
Society and Natural Resources, 15: 949-957

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This article examines the development and implementation of a grass-roots elephant conservation program based upon the Samburu people’s perceptions and knowledge of elephants in the areas surrounding the Samburu and Buffalo Springs National Reserves in northern Kenya. Ethnographic methods were used to understand these perceptions and demonstrated that strong customs and traditions for conserving wildlife, particularly elephants, exist among the Samburu people. It became evident that these customs are changing, given various factors influencing Samburu culture and younger generations. The use of economic incentives is a widely accepted method to foster positive attitudes and behavior toward wildlife. The value of using ethnographic methods to reinforce positive indigenous knowledge about wildlife, however, is underestimated.

Lala, F., Chiyo, P.I., Omondi, P., Okita-Ouma, B., Kanga, E., Koskei, M., Tiller, L., Morris, A.W., Severud, W.J. and Bump, J.K., 2022
Influence of infrastructure, ecology, and underpass-dimensions on multi-year use of Standard Gauge Railway underpasses by mammals in Tsavo, Kenya. (2022)
Scientific reports, 12(1), pp.1-14.

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Rail and road infrastructure is essential for economic growth and development but can cause a gradual loss in biodiversity and degradation of ecosystem function and services. We assessed the infuence of underpass dimensions, fencing, proximity to water and roads, Normalized Diference Vegetation Index (NDVI), presence of other species and livestock on underpass use by large and medium-sized mammals. Results revealed hyenas and leopards used the underpasses more than expected whereas girafes and antelopes used the underpasses less than expected. Generalized linear mixed-efects models revealed that underpass height infuenced use by wildlife, with several species preferring to use taller underpasses. Electric fencing increased underpass use by funneling species towards underpasses, except for elephants and black-backed jackal for which it reduced underpass passage. We also found that the use of underpasses by livestock reduced the probability of use by nearly 50% for wildlife species. Carnivore species were more likely to cross underpasses used by their prey. Bufalo, livestock, and hyenas used underpasses characterized by vegetation with higher NDVI and near water sources while baboons, dik-diks and antelope avoided underpasses with high NDVI. Our fndings suggest a need for diverse and comprehensive approaches for mitigating the negative impacts of rail on African wildlife.

Lamprey, R., Ochanda, D., Brett, R., Tumwesigye, C., Douglas-Hamilton, I.
Cameras replace human observers in multi-species aerial counts in Murchison Falls, Uganda (2020)
Remote Sensing in Ecology and Conservation published by John Wiley & Sons Ltd on behalf of Zoological Society of London https://zslpublications.onlinelibrary.wiley.com/doi/full/10.1002/rse2.154 doi: 10.1002/rse2.154

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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.

Lamprey, R., Pope, F., Ngene, S., Norton-Griffith, M., Frederick, H., Okita-Ouma, B., Douglas-Hamilton, I.
Comparing an automated high-definition oblique camera system to rear-seat-observers in a wildlife survey in Tsavo, Kenya: Taking multi-species aerial counts to the next level (2019)
Biological Conservation https://doi.org/10.1016/j.biocon.2019.108243

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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.

Litoroh, M., Ihwagi, F.I., Mayienda, R., Bernard,J., Douglas-Hamilton, I.
Total Aerial Count of Elephants in Laikipia-Samburu Ecosystem in November 2008. (2010)
Kenya Wildlife Service

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The total aerial count of Elephants in Laikipia-Samburu Ecosystem and for the second time in Marsabit was conducted between 24th and 28th November 2008. Total counts of elephants, their carcasses and Gravy’s Zebra were done. Livestock and other wildlife species mainly the large herbivores were also counted. Coming six years after the preceding total count of Omondi et. al. 2002, the data provides a basis for assessment of the trends in the ecosystem. Within Laikipia-Samburu ecosystem, 7415 elephants were counted, up from 5447 elephants counted in 2002, an estimated annual growth rate of 5.3%. West Isiolo/Samburu block hosted 26% of the elephant population in Ewaso Ecosystem at the highest density of 1.332 elephants per square kilometer. Most of these elephants were encountered in Buffalo Springs National Reserve. Ninety five elephant carcasses were recorded.

Maisels, F. et al
Devastating Decline of Forest Elephants in Central Africa. (2013)
PLOS ONE 10.1371/journal.pone.0059469

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African forest elephants – taxonomically and functionally unique–are being poached at accelerating rates, but we lack range-wide information on the repercussions. Analysis of the largest survey dataset ever assembled for forest elephants (80 foot-surveys; covering 13,000 km; 91,600 person-days of fieldwork) revealed that population size declined by ca. 62% between 2002–2011, and the taxon lost 30% of its geographical range. The population is now less than 10% of its potential size, occupying less than 25% of its potential range. High human population density, hunting intensity, absence of law enforcement, poor governance, and proximity to expanding infrastructure are the strongest predictors of decline. To save the remaining African forest elephants, illegal poaching for ivory and encroachment into core elephant habitat must be stopped.

Marshal, J.P., Azhar Rajah, A., Parrini, F., Henley, M., Henley, S.R., Erasmus, B.F.N.
Scale-dependent selection of greenness by African elephants in the Kruger-private reserve transboundary region, South Africa. (2010)
European Journal of Wildlife Research DOI: 10.1007/s10344-010-0462-1

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Foraging behaviour and habitat selection occur as hierarchical processes. Understanding the factors that govern foraging and habitat selection thus requires investigation of those processes over the scales at which they occur. We investigated patterns of habitat use by African elephants (Loxodonta africana) in relation to vegetation greenness to investigate the scale at which that landscape attribute was most closely related to distribution of elephant locations. We analysed Global Positioning System radio-collar locations for 15 individuals, using the Normalized Difference Vegetation Index as a representation of vegetation greenness in a Geographic Information Systems framework. We compared the importance of vegetation greenness at three spatial scales: the total home range, the seasonal home range and the 16-day home range.

Martin, E., Stiles, D.
The Ivory Markets of Europe. (2005)
Save The Elephants

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This report is the fourth in a series of surveys that depict the status and trends of the elephant ivory markets in a particular region of the world. Previous surveys covered Africa (Martin and Stiles 2000), South and South East Asia (Martin and Stiles 2002) and East Asia (Martin and Stiles 2003). This report deals with five countries in Europe: Germany, the UK, France, Spain and Italy, in relative order of market scale. These countries were selected on the basis of the size of their economies, and thus buying power, and on informants’ reports in Africa and Asia of the principal European buyers of worked ivory in their regions. The surveys were carried out between April and November 2004, and the work was sponsored by Save the Elephants and Care for the Wild International.

Martin, E., Stiles, D.
The Ivory Markets in the USA (2008) Save the Elephants & Care for the Wild. (2008)
Save the Elephants & Care for the Wild  

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This report is the fifth in a series of surveys that depict the status and trends of the elephant ivory markets in a?particular region of the world. This investigation covered the United States of America (USA) and Vancouver,?Canada. It differs significantly from previous studies of this trade by quantifying the nature and scale of the?market. The investigators made the survey between March and December 2006 and March and May 2007.?Seventeen cities and towns were selected for study based on their population size and wealth, and tourist?importance.

Martin, E., Stiles, D.
The Ivory Markets of Africa (2002)
Save The Elephants.

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Although the international trade in elephant ivory is banned by almost every country, there are significant quantities of tusks and carved items moving illicitly within Africa, especially in Central and West Africa, and from the continent to markets in eastern Asia. Within some African countries there is also a large internal (domestic) trade in ivory objects. ??Unfortunately, prior to this study, there have been few statistics and little information available on the present-day ivory trade markets in Africa. With the CITES approved one-off sales of government ivory stocks in Namibia, Botswana and Zimbabwe in April 1999, it has become even more important to have reliable data on the prices and quantities of raw and worked ivory in the principal markets of Africa.

Martin, E., Stiles, D.
The South and South East Asian Ivory Markets. (2002)
Save The Elephants

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The purpose of this report is first to present data on the current status of the ivory trade in the major markets of South and South East Asia. These data are needed by relevant government authorities, international and national wildlife conservation organizations and by the Convention on the International Trade in Endangered Species of Wild Fauna and Flora (CITES) in order to comprehend the scale of the ivory market in the respective countries today. ??The second objective is to set a base line of ivory trade indicators from which to assess what is happening to the trade so that future monitoring and evaluation can be carried out on the effectiveness of policies, laws and enforcement activities relating to the internal and international trade in ivory.

Martin, E., Stiles, D.
The Ivory Markets of East Asia. (2003)
Save The Elephants.

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This report presents the results of the third in a series of surveys that describe the status and trends of the elephant ivory trade in a region of the world, in this case East Asia. The previous reports covered Africa (Martin and Stiles 2000; Stiles and Martin 2001) and South and South East Asia (Martin and Stiles 2002; Stiles and Martin 2002). ??The places surveyed for this report were Japan, Hong Kong, China, Taiwan, and South Korea. The purpose of these ivory trade surveys is to gather data on a set of indicators that portray the scale of the market in ivory in order that governments, wildlife conservation organizations and CITES representatives can appreciate the extent to which ivory is traded in selected countries.

Martin, E., Vigne, L.
Hong Kong’s Ivory – More items for sale than in any other city in the world. (2015)
Save The Elephants

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Hong Kong displays for retail sale more elephant and mammoth ivory items than any other city in the world surveyed for ivory. There are also large, intricately carved pieces of mammoth ivory that can be legally taken out of Hong Kong, as they were carved from the extinct mammoth. This is in contrast to mainly trinkets, especially jewellery, that are carved from elephant ivory. They are illegal to export, but being small, these items are easily smuggled out in personal luggage. Hong Kong is one of the biggest tourist destinations in Asia. The city has a population of about 7 million people yet attracts over 50 million visitors a year, mostly mainland Chinese. They come to shop, including for luxury products such as ivory because these items are less expensive than on the mainland.

Michelmore, F., Beardsley, K., Barnes, R., Douglas-Hamilton, I.
A model illustrating the changes in forest elephant numbers caused by poaching. (1994)
African Journal of Ecology.

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A geographic information system (GIS) was used to analyse field data on the abundance of elephant dung-piles. For each country, the GIS was used to create contours representing distances from roads or rivers. The area of forest between each contour was then calculated. The curvilinear relationships between dung-pile density and distance to the nearest road or village were then used to calculate the numbers of dung-piles between contours and the total for each country. Comparisons between undisturbed and heavily poached elephant populations suggest that the total forest elephant population in central Africa has been reduced by 44% as a result of ivory poaching. Forest elephants may be more vulnerable to poaching than previously thought because, for example, two-thirds of Congo's elephants live within two day's walk of a road or navigable river.

Mortimer B., Walker J. A., Lolchuragi D. S., Reinwald M., Daballen D
Noise matters: elephants show risk-avoidance behaviour in response to human-generated seismic cues (2021)
Proc. R. Soc. B.2882021077420210774 http://doi.org/10.1098/rspb.2021.0774

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Abstract African elephants (Loxodonta africana) use many sensory modes to gather information about their environment, including the detection of seismic, or ground-based, vibrations. Seismic information is known to include elephant-generated signals, but also potentially encompasses biotic cues that are commonly referred to as ‘noise’. To investigate seismic information transfer in elephants beyond communication, here we tested the hypothesis that wild elephants detect and discriminate between seismic vibrations that differ in their noise types, whether elephant- or human-generated. We played three types of seismic vibrations to elephants: seismic recordings of elephants (elephant-generated), white noise (human-generated) and a combined track (elephant- and human-generated). We found evidence of both detection of seismic noise and discrimination between the two treatments containing human-generated noise. In particular, we found evidence of retreat behaviour, where seismic tracks with human-generated noise caused elephants to move further away from the trial location. We conclude that seismic noise are cues that contain biologically relevant information for elephants that they can associate with risk. This expands our understanding of how elephants use seismic information, with implications for elephant sensory ecology and conservation management.

Mortimer, B., Rees, W. L., Koelemeijer, P, and Nissen-Meyer, T.
Classifying elephant behaviour through seismic vibrations (2018)
Current Biology. Volume 28, Issue 9, 7 May 2018, Pages R547-R548 https://doi.org/10.1016/j.cub.2018.03.062

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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.

Mugo, G., Tiller, L. and King, L.
Mapping Floristic Composition Using Sentinel-2A and a Case Study Evaluation of Its Application in Elephant Movement Ecology in Sagalla, Kenya (2022)
Remote Sensing, 14(21), p.5386.

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Abstract The quantification of vegetation structure and composition at local and global scales provides valuable information for understanding the balance of the natural and human-made environment, which is crucial for natural resource planning and management, and the sustenance of ecosystem biodiversity. In this study, we proposed using the Sentinel 2A imagery to classify vegetation cover into communities based on the floristic association of individual vegetation species. We apply traditional remote sensing techniques to process the satellite image and identify training regions of interest (ROI) which are thoroughly assessed for spectral uniqueness before using the pixel-based supervised classification algorithms for our classification. Ground truthing assessment and species dominance computations are done to determine the vegetation community composition and naming based on floristic associations. We apply the floristic compositions output in analysing elephant movement tracks in the area, to assess the potential influence the location of specific vegetation species and communities utilized by elephants has on their movement and presence, as well as on elephant bulls and family groupings. The results show that the 10 m spatial resolution Sentinel-2A is suitable for investigating and mapping vegetation species in communities for large-scale mapping operations. We determined Near-Infrared band 8 and shortwave Infrared band 11 as key for identifying and differentiating ROIs at the floristic association community vegetation mapping level. We attained an overall accuracy of 87.395%. The analysis proved the 10 m spatial resolution of Sentinel 2A to be sufficient in distinguishing vegetation communities, including those with similar dominant species but variations in other contributing species. We also found a direct connection between vegetation location and elephant movement based on the summative analysis of utilised vegetation by the different elephant groupings. Bull elephants were predominantly present in areas with Combretum, family groups in areas with Commiphora, and mixed groups with both bulls and families in areas with Commiphora, and Cissus. This study shows the value that remote-sensing scientific support can offer conservationists and governments in objective evidence-based land management, policy making and governance.

Ngene, S., Njumbi, S., Nzisa, M., Kimitei, K., Mukeka, J., Muya, S., Ihwagi, F., Omondi, P.
Status and trends of the elephant population in the Tsavo–Mkomazi ecosystem (2013)
Pachyderm No. 53 January–June 2013

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This paper updates the data on the population status of elephants in the Tsavo–Mkomazi ecosystem. Data were acquired through aerial census of elephants in the ecosystem, from 7 to 12 February 2011. The census covered an area approximately 48,319 km2, which was divided into 44 counting blocks. Each block was assigned to a specific aircraft; the crew consisted of a pilot, front-seat observer and two rear-seat observers for the four-seater light aircraft, and a pilot and an observer for a two-seater light aircraft. The census lasted five days and involved nine light aircraft and about 252 hours of actual counting time, representing a mean search rate of about 191 km2/hr. A total of 12,573 elephants were counted, indicating a modest increase of 2% after the 2008 census and a 96% increase after the 1988 census (n = 6,399).

Ogola, P.
Demographic Status of the Meru Elephant Population. (2003)
Final Report to the Elephant Research Fund/KWS and Save the Elephants

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406 elephant were encountered on the ground during a one year (Sept 2001-Sept 2002) study focused on identifying individuals and studying the population dynamics of elephant in Meru National Park. (This number did not include 17 animals recently transferred from Sweetwaters and another 22 seen from the air only) The population exhibited a seasonal migration pattern to areas north and northwest of the Park which may put some animals as risk to due to the general insecurity of the area. The demographic data collected suggest the population remained static in the early 1990s but experienced a high rate of growth in the last few years. This increase since the late 1990s was attributed to: translocations of over 70 elephant into the Park, high calf recruitment (average 5%) since 1997, and low adult mortality compared with that of the previous decade.

Ojwang, G.O., Wargute, P.W., Said, M.Y., Worden, J.S., Davidson, Z., Muruthi, P., Kanga, E., Ihwagi, F., Okita-Ouma, B.
Wildlife Migratory Corridors and Dispersal Areas: Kenya Rangelands and Coastal Terrestrial Ecosystems (2017)
Copyright © 2017 Government of the Republic of Kenya

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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.

Okello, J.B.A., Masembe, C., Rasmussen, H.B., Wittemyer, G., Omondi, P., Kahindi, O., Muwanika, V.B., Arctander, P., Douglas-Hamilton, I., Nyakaana, S., Siegismund, H.R.
Population Genetic Structure of Savannah Elephants in Kenya: Conservation and Management Implications (2008)
Journal of Heredity 2005:96(6):679–687

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We investigated population genetic structure and regional differentiation among African savannah elephants in Kenya using mitochondrial and microsatellite markers. We observed mitochondrial DNA (mtDNA) nucleotide diversity of 1.68% and microsatellite variation in terms of average number of alleles, expected and observed heterozygosities in the total study population of 10.20, 0.75, and 0.69, respectively. Hierarchical analysis of molecular variance of mtDNA variation revealed significant differentiation among the 3 geographical regions studied (FCT 5 0.264; P , 0.05) and a relatively lower differentiation among populations within regions (FSC 5 0.218; P , 0.0001). Microsatellite variation significantly?differentiated among populations within regions (FSC 5 0.019; P , 0.0001) but not at the regional levels (FCT 5 0.000; P . 0.500).

Okello, J.B.A., Wittemyer, G., Rasmussen, H.B., Arctander, P., Nyakaana, S., Douglas-Hamilton, I., Siegismund, H.R.
Effective population size dynamics reveal impacts of historic climatic events and recent anthropogenic pressure in African elephants. (2008)
Molecular Ecology

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Two hundred years of elephant hunting for ivory, peaking in 1970–1980s, caused local extirpations and massive population declines across Africa. The resulting genetic impacts on surviving populations have not been studied, despite the importance of understanding the evolutionary repercussions of such human-mediated events on this keystone species. Using Bayesian coalescent-based genetic methods to evaluate time-specific changes in effective population size, we analysed genetic variation in 20 highly polymorphic microsatellite loci from 400 elephants inhabiting the greater Samburu-Laikipia region of northern Kenya. This area experienced a decline of between 80% and 90% in the last few decades when ivory harvesting was rampant.

Okello, J.B.A., Wittemyer, G., Rasmussen, H.B., Douglas-Hamilton, I., Nyakaana, S., Arctander, P.S.
Noninvasive Genotyping and Mendelian Analysis of Microsatellites in African Savannah Elephants. (2005)
Journal of Heredity 2005:96(6)

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We obtained fresh dung samples from 202 (133 mother-offspring pairs) savannah elephants (Loxodonta africana) in Samburu, Kenya, and genotyped them at 20 microsatellite loci to assess genotyping success and errors. A total of 98.6% consensus genotypes was successfully obtained, with allelic dropout and false allele rates at 1.6% (n 5 46) and 0.9% (n 5 37) of heterozygous and total consensus genotypes, respectively, and an overall genotyping error rate of 2.5% based on repeat typing. Mendelian analysis revealed consistent inheritance in all but 38 allelic pairs from mother-offspring, giving an average mismatch error rate of 2.06%, a possible result of null alleles, mutations, genotyping errors, or inaccuracy in maternity assignment.

Okita-­Ouma B., Lala F., Koskei M., Mwazo A., Kibara D., King L., Kanga E. & Douglas-Hamilton I.
Movements of satellite-­linked collared elephants and other wildlife in relation to the Standard Gauge Railway (SGR) and highways in Tsavo ecosystem, Kenya (2016)
Save The Elephants and Kenya Wildlife Service, Nairobi

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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.

Okita-Ouma B., Lala F., Koskei M., Mwazo A., Kibara D., King L., & Douglas-Hamilton I.
Tracking and monitoring of elephant movements along the Standard Gauge Railway and highways in the Tsavo Ecosystem, Kenya (March 2016 – June 2017) (2017)
Save The Elephants and Kenya Wildlife Service. Typescript 28 pages

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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.

Okita-Ouma, B., Koskei, M., Tiller, L., Lala, F., King, L., Moller, R., Amin, R. and Douglas-Hamilton, I.
Effectiveness of wildlife underpasses and culverts in connecting elephant habitats: a case study of new railway through Kenya’s Tsavo National Parks (2021)
Afr J Ecol. https://doi.org/10.1111/aje.12873

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Abstract Transportation networks can be a major impediment to wildlife movements. We assessed the use of wildlife underpasses and culverts along a newly constructed railway in Kenya's Tsavo National Parks by African elephants (L. africana). We collared ten elephants with GPS satellite transmitters within 20 km of the railway in March 2016 and analysed their movement data to March 2019. Eight elephants used the underpasses although one did not cross the adjacent highway. The remaining two neither used the underpasses nor crossed the highway despite ranging in the vicinity. Their median speed significantly increased to 0.65 km/hr from 0.45 km/hr before crossing the railway, then slowed to 0.32 km/hr after crossing. Females in family groups moved faster than the lone bulls when using the underpasses. Seventy-eight per cent of all crossings made were at night. The fast speeds and the nocturnal patterns are behavioural responses of elephants in risky landscapes or under stress. Disturbance from vehicles traffic on the adjacent highway and from newly developed human settlements may have limited use of underpasses. Wildlife crossing structures, signage and speed bumps along the highway; relocation of the illegal human settlements; and inter-agency coordination are requisites for enhancing Tsavos' elephant habitat connectivity.

Okita-Ouma, B., Lala, F., Moller, R., Koskei, M., Kiambi, S., Daballen, D., Leadismo, C., Mijele, D., Poghon, J., King, L., Pope, F., Wittemyer, G., Wall, J., Goss, S., Obrien, R., Douglas-Hamilton, I.
Preliminary indications of the effect of infrastructure development on ecosystem connectivity in Tsavo National Parks, Kenya (2016)
Pachyderm No. 57 July 2015–June 2016

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Balancing conservation and infrastructure developmentConserving land and ecosystem connectivity for wildlife is increasingly a global challenge as demand for infrastructure development to meet growing human population needs encroaches in many traditional wildlife areas. The survival of wildlife species in arid and semi-arid systems requires interconnected landscapes, and limiting animal movement greatly reduces the system’s ability to sustain viable wildlife populations (Vasudev et al. 2015). Major infrastructural developments such as multi-lane highways and railways can sever wildlife movement often with negative consequences (Clevenger and Waltho 2005; van-der-Ree et al. 2011; Xia et al. 2007).

Olindo, P.M., Douglas-Hamilton, I., Hamilton, P.
The 1988 Tsavo Elephant Count. (1988)
Kenya Wildlife Conservation and Management Department

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Results from a large scale aerial survey of large mammals in the Tsavo ecosystems in February are compared to earlier surveys. 4,327 elephants were counted in the Park, a decline of 75% since the 1972 count. Another 1,036 were counted in surrounding areas, a decline of 87% over the same period. Results confirm the downward trend noted in KREMU sample counts which suggest that the Tsavo ecosystem 1969 population of 42,000 elephants stood at about 5,700 in 1987. The worse affected areas were Mkomazi GR where numbers are down by 96% and Galana Ranch where they have fallen by 98%. In Taita-Taveta District area near the Park they decreased by 31% while near the Taita Hills and Salt Lick Lodges an increase in numbers suggests that tourism may offer some degree of protection from poaching.

Omondi P., Bitok E., Kahindi O., Mayienda, R.
Total Aerial Count of Elephants in Samburu-Laikipia EcoSystem: June 2002. (2002)
Kenya Wildlife Service

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The total aerial count of elephants in Laikipia/Samburu ecosystem was carried out between 20th and 24th of June 2002, During the census, total counts of elephants, elephant carcasses and buffaloes was done. Livestock numbers (cattle and shoats) were estimated. As a MIKE site the count provided baseline data for monitoring poaching levels and elephant trends in the ecosystem. A total of 5,447elephants were counted during the survey (2,206 or 40.5% in Samburu and 3,241 or 59.5% in Laikipia).  The overall increase since 1999 in the entire ecosystem was 58.5%. The number of carcasses counted was 64 with only one being fresh and three recent while the rest were old or very old. The carcass ratio for the ecosystem was 1.16% - a decline as compared to 1999 (2.8%), however the percentage of recent carcasses rose from 6% to 6.25% (1999 and 2002 respectively).

Padariya, M., Jooste, M.L., Hupp, T., Fåhraeus, R., Vojtesek, B., Vollrath, F., Kalathiya, U. and Karakostis, K.
The Elephant evolved p53 isoforms that escape mdm2-mediated repression and cancer (2022)
Molecular biology and evolution, 39(7), p.msac149.

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Abstract The p53 tumor suppressor is a transcription factor with roles in cell development, apoptosis, oncogenesis, aging, and homeostasis in response to stresses and infections. p53 is tightly regulated by the MDM2 E3 ubiquitin ligase. The p53–MDM2 pathway has coevolved, with MDM2 remaining largely conserved, whereas the TP53 gene morphed into various isoforms. Studies on prevertebrate ancestral homologs revealed the transition from an environmentally induced mechanism activating p53 to a tightly regulated system involving cell signaling. The evolution of this mechanism depends on structural changes in the interacting protein motifs. Elephants such as Loxodonta africana constitute ideal models to investigate this coevolution as they are large and long-living as well as having 20 copies of TP53 isoformic sequences expressing a variety of BOX-I MDM2-binding motifs. Collectively, these isoforms would enhance sensitivity to cellular stresses, such as DNA damage, presumably accounting for strong cancer defenses and other adaptations favoring healthy aging. Here we investigate the molecular evolution of the p53–MDM2 system by combining in silico modeling and in vitro assays to explore structural and functional aspects of p53 isoforms retaining the MDM2 interaction, whereas forming distinct pools of cell signaling. The methodology used demonstrates, for the first time that in silico docking simulations can be used to explore functional aspects of elephant p53 isoforms. Our observations elucidate structural and mechanistic aspects of p53 regulation, facilitate understanding of complex cell signaling, and suggest testable hypotheses of p53 evolution referencing Peto’s Paradox.

Parker, Ian S. C
The ivory trade. (1979)
Report to USFWS

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A multi volume typescript report to Iain Douglas-Hamilton to be used as part of his final 1979 report to US Fish and Wildlife Service, Washington, D. C. Focused on the ivory trade in terms of size with respect to utilization of tusks as well as economic base of the trade this consultancy provides data on the history and present ivory legal and illegal trade within Africa and between Africa and major importing nations.

Parker, J.M., Brown, J.L., Hobbs, N.T., Boisseau, N.P., Letitiya, D., Douglas-Hamilton, I. and Wittemyer, G.
Social support correlates with glucocorticoid concentrations in wild African elephant orphans. (2022)
Communications Biology, 5(1), pp.1-9.

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Abstract Social relationships have physiological impacts. Here, we investigate whether loss of the mother/offspring relationship has lasting effects on fecal glucocorticoid metabolite (fGCM) concentrations in wild African elephant orphans several years following their mothers’ deaths. We find no difference in fGCM concentrations between orphans and nonorphans, but find lower fGCM concentrations in elephants with more age mates in their family. We also unexpectedly identify lower concentrations in orphans without their natal family versus nonorphans and natal orphans, which we speculate may be due to the development of hypocortisolism following a prolonged period without familial support. An index of plant productivity (i.e. food) shows the largest correlation with fGCM concentrations. Our findings indicate no lasting differences in glucocorticoid concentrations of surviving orphan elephants who are with their family, suggest the presence of age mates may reduce glucocorticoid concentrations in elephants, and emphasize that basic survival needs are the primary regulators of the stress response.

Parker, J.M., Webb, C.T., Daballen, D., Goldenberg, S.Z., Lepirei, J., Letitiya, D., Lolchuragi, D., Leadismo, C., Douglas-Hamilton, I., Wittemyer, G.
Poaching of African elephants indirectly decreases population growth through lowered orphan survival (2021)
Current Biology, 2021, ISSN 0960-9822, https://doi.org/10.1016/j.cub.2021.06.091. (https://www.sciencedirect.com/science/article/pii/S0960982221009209)

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Prolonged maternal care is vital to the well-being of many long-lived mammals.1 The premature loss of maternal care, i.e., orphaning, can reduce offspring survival even after weaning is complete.2, 3, 4, 5 However, ecologists have not explicitly assessed how orphaning impacts population growth. We examined the impact of orphaning on population growth in a free-ranging African elephant population, using 19 years of individual-based demographic monitoring data. We compared orphan and nonorphan survival, performed a sensitivity analysis to understand how population growth responds to the probability of being orphaned and orphan survival, and investigated how sensitivity to these orphan parameters changed with level of poaching. Orphans were found to have lower survival compared to nonorphaned age mates, and population growth rate was negatively correlated with orphaning probability and positively correlated with orphan survival. This demonstrates that, in addition to its direct effects, adult elephant death indirectly decreases population growth through orphaning. Population growth rate’s sensitivity to orphan survival increased for the analysis parameterized using only data from years of more poaching, indicating orphan survival is more important for population growth as orphaning increases. We conclude that orphaning substantively decreases population growth for elephants and should not be overlooked when quantifying the impacts of poaching. Moreover, we conclude that population models characterizing systems with extensive parental care benefit from explicitly incorporating orphan stages and encourage research into quantifying effects of orphaning in other social mammals of conservation concern.

Polansky, L., Douglas-Hamilton, I., Wittemyer, G.
Using diel movement behavior to infer foraging strategies related to ecological and social factors in elephants. (2013)
Movement Ecology 2013, 1:13 doi:10.1186/2051-3933-1-13

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Abstract Adaptive movement behaviors allow individuals to respond to fluctuations in resource quality and distribution in order to maintain fitness. Classically, studies of the interaction between ecological conditions and movement behavior have focused on such metrics as travel distance, velocity, home range size or patch occupancy time as the salient metrics of behavior. Driven by the emergence of very regular high frequency data, more recently the importance of interpreting the autocorrelation structure of movement as a behavioral metric has become apparent. Studying movement of a free ranging African savannah elephant population, we evaluated how two movement metrics, diel displacement (DD) and movement predictability (MP - the degree of autocorrelated movement activity at diel time scales), changed in response to variation in resource availability as measured by the Normalized Difference Vegetation Index. We were able to capitalize on long term (multi-year) yet high resolution (hourly) global positioning system tracking datasets, the sample size of which allows robust analysis of complex models. We use optimal foraging theory predictions as a framework to interpret our results, in particular contrasting the behaviors across changes in social rank and resource availability to infer which movement behaviors at diel time scales may be optimal in this highly social species. http://www.movementecologyjournal.com/content/1/1/13

Polansky, L., Wittemyer, G.
A framework for understanding the architecture of collective movements using pairwise analyses of animal movement data. (2010)
Journal of Royal Society Interface. doi:10.1098/rsif.2010.0389

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The study of collective or group-level movement patterns can provide insight regarding the socio-ecological interface, the evolution of self-organization and mechanisms of inter-individual information exchange. The suite of drivers influencing coordinated movement trajectories occur across scales, resulting from regular annual, seasonal and circadian stimuli and irregular intra- or interspecific interactions and environmental encounters acting on individuals. Here, we promote a conceptual framework with an associated statistical machinery to quantify the type and degree of synchrony, spanning absence to complete, in pairwise movements. The application of this framework offers a foundation for detailed understanding of collective movement patterns and causes.

Polansky, L., Wittemyer, G., Cross, P.C., Tambling, C.J., Getz, W.M.
From moonlight to movement and synchronized randomness: Fourier and wavelet analyses of animal location time series data. (2010)
Ecology, 91(5), 2010, pp. 1506–1518

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High-resolution animal location data are increasingly available, requiring analytical approaches and statistical tools that can accommodate the temporal structure and transient dynamics (non-stationarity) inherent in natural systems. Traditional analyses often assume uncorrelated or weakly correlated temporal structure in the velocity (net displacement) time series constructed using sequential location data. We propose that frequency and time– frequency domain methods, embodied by Fourier and wavelet transforms, can serve as useful probes in early investigations of animal movement data, stimulating new ecological insight and questions. We introduce a novel movement model with time-varying parameters to study these methods in an animal movement context.

Raizman, E.A., Rasmussen, H.B., King, L.E., Ihwagi, F.W., Douglas-Hamilton, I.
Feasibility study on the spatial and temporal movement of Samburu’s cattle and wildlife in Kenya using GPS radio-tracking, remote sensing and GIS. (2013)
PREVET (2013) http://dx.doi.org/10.1016/j.prevetmed.2013.04.007

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The study was conducted to assess the technical feasibility of studying the spatial and temporal interaction of traditionally herded livestock and wildlife using global positioning system (GPS) tracking technology in Northern Kenya. Two types of collars were used on nine cows: radio frequency and global system for mobile communications (GSM) collars and GPS-satellite (SAT) collars. Full results of cattle tracking were available for eight cows (3 GSM and 5 SAT) tracked between July 2008 and September 2010. A cumulative total of 1556 tracking days was recorded over the 17 month period. On average cows walked 10,203 m/day (average total monthly distance walked was 234 km). Significant seasonal differences were found; on average cows walked 9.607 m and 10,392 m per day in the rainy and the dry seasons, respectively.

Rasmussen, H.B.
Reproductive tactics of male African savannah elephants (Loxodonta africana) (2005)
Thesis submitted in fulfilment of the requirements for the degree of Doctor of Philosophy at the University of Oxford Michaelmas Term 2005

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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.

Rasmussen, H.B.
Mating strategies in the Male African Elephant, Loxodonta africana. (2001)
Masters thesis, Aarhus Univeristy, Denmark

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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.

Rasmussen, H.B., Ganswindt, A., Douglas-Hamilton, I., Vollrath, F.
Endocrine and behavioral changes in male African elephants: Linking hormone changes to sexual state and reproductive tactics. (2008)
Elsevier. Hormones and Behavior

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Hormones play a crucial role in mediating genetic and environmental effects into morphological and behavioral phenotypes. In systems with alternative reproductive tactics (ART) shifts between tactics are hypothesized to be under proximate hormonal control. Most studies of the underlying endocrine changes behind ART have focused on !sh and amphibians rather than mammals and few have investigated the potential interaction between different endocrine axes in regulating shifts between conditional dependent tactics. Using a combination of endocrine and behavioral data from male African elephants we expand on our previously published analysis and show that the initial increase in androgens predates the behavioral shifts associated with reproductively active periods, supporting the role of androgens in activating sexually active periods in males.

Rasmussen, H.B., Kahindi O., Vollrath, F., Douglas-Hamilton, I.
Estimating elephant densities from wells and droppings in dried out riverbeds. (2005)
African Journal of Ecology , Vol. 43 Issue 4 Page 312 December 2005

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In this paper, we present a new method for estimating elephant densities by counting elephant wells and dung boli within dry seasonal ?ooding rivers. A combination of aerial and ground counts of elephant wells and dung boli in the Ewaso Ngiro River were related to elephant numbers, obtained from an on-going monitoring program of individually identi?ed elephants in Samburu and Buffalo Spring National Reserves, Kenya. The total number of elephant observations was highly correlated with both densities of wells and dung boli at a spatial resolution of 4-km river-section. This indicates that both wells and droppings can be used for estimating relative densities at such spatial resolution.

Rasmussen, H.B., Okello,J.B.A., Wittemyer, G., Siegismund,H.R., Arctander,P., Vollrath,F., and Douglas-Hamilton, I.
Age – and tactic-related paternity success in male African elephants. (2007)
Rasmussen, H.B., Okello,J.B.A., Wittemyer, G., Siegismund,H.R., Arctander,P., Vollrath,F., and Douglas-Hamilton, I. ?Age- and tactic-related paternity success in male African elephants (2007) ?Behavioral Ecology

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Information on age- and tactic-related paternity success is essential for understanding the lifetime reproductive strategy of males and constitutes an important component of the fitness trade-offs that shape the life-history traits of a species. The degree of reproductive skew impacts the genetic structure of a population and should be considered when developing conservation strategies for threatened species. The behavior and genetic structure of species with large reproductive skew may be disproportionately impacted by anthropogenic actions affecting reproductively dominant individuals. Our results on age- and tactic-specific paternity success in male African elephants are the first from a free-ranging population and demonstrate that paternity success increases dramatically with age, with the small number of older bulls in the competitive state of musth being the most successful sires.

Rasmussen, H.B., Wittemyer, G., Douglas-Hamilton, I.
Estimating age of immobilized elephants from teeth impressions using dental silicon. (2005)
African Journal of Ecology 43:215-219

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High precision condensation dental silicon, ZetalaborTM, was used to create moulds of the lower jaw molars from 22 immobilized African elephants (Loxodonta africana Blumenback) during radio collaring operations. These moulds were used to determine the elephant’s age using Laws and Jachmann’s molar aging criteria. The technique proved easy and fast and produced useful imprints in 90% of the cases. We found our age estimates, based on physical appearance, made prior to immobilizations were relatively accurate, with 75% within ±3 years and 95% within ±5 years from the age indicated from molar evaluation. When re-collaring the same individuals in 2–3 years, new moulds will be made to compare a known time period with the degree of tooth wear. This will provide verification of Laws age estimates from free-ranging elephants.

Rasmussen, H.B., Wittemyer, G., Douglas-Hamilton, I.
Predicting time-specific changes in demographic processes using remote-sensing data. (2006)
Journal of Applied Ecology (2006) 43, 366–376

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1. Models of wildlife population dynamics are crucial for sustainable utilization and?management strategies. Fluctuating ecological conditions are often key factors influencing?both carrying capacity, mortality and reproductive rates in ungulates. To be reliable,?demographic models should preferably rely on easily obtainable variables that are?directly linked to the ecological processes regulating a population.?2. We compared the explanatory power of rainfall, a commonly used proxy for variability?in ecological conditions, with normalized differential vegetation index (NDVI), a?remote-sensing index value that is a more direct measure of vegetation productivity, to?predict time-specific conception rates of an elephant population in northern Kenya.?Season-specific conception rates were correlated with both quality measures.

Rasmussen, L.E.L., Wittemyer, G.
Chemosignaling of musth by individual wild African elephants, (Loxodonta africana): implications for conservation and management. (2002)
Proceedings of the Royal Society: Biological Sciences 269:853-860

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Elephants have extraordinary olfactory receptive equipment, yet this sensory system has been only minimally?investigated in wild elephants. We present an in-depth study of urinary chemical signals emitted?by individual, behaviourally characterized, wild male African elephants, investigating whether these compounds ?were the same, accentuated, or diminished in comparison with captive individuals. Remarkably,?most emitted chemicals were similar in captive and wild elephants with an exception traced to drought induced? dietary cyanates among wild males. We observed developmental changes predominated by the ?transition from acids and esters emitted by young males to alcohols and ketones released by older males.?

Roth, H.H., Douglas-Hamilton, I.
Distribution and status of elephants in West Africa. (1991)
Mammalia Vol 55 Issue 4 p489-527;table;figs;photos;maps;refs.

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The taxonomy of West African elephants is reviewed along with their distribution during 1976-1984 study done in a biogeographical context. At that time they inhabited roughly 232,000 sq km, 6-7% of the area they had occupied at the turn of the century. The diminution of elephant range was greatest in the Sahel and Guinean savannah zone of which only about 2.8% and 2.3% respectively are still inhabited by elephants as compared to 6.7% in the Sudanian and 16% in the Guinean forest zone. Reasons for the decline of elephants in West Africa are analyzed. Accepting inaccuracies of elephant censusing, surviving elephants in West Africa were estimated at roughly 17,000 of which about 5,200 were considered to be forest elephants. Of these only about 11,500 (3,500 forest and 8,000 savannah elephants) live in or around national parks and game reserves. Conservation and chances for survival of elephants are discus

Ruto, E.
ASSESSING THE POTENTIAL IMPACT OF CLIMATE VARIABILITY AND ANTHROPOLOGICAL ACTIVITIES ON HONEY BEE FODDER PLANTS IN SAGALLA, TAITA TAVETA COUNTY, KENYA (2018)
MSc. Thesis SUBMITTED IN PARTIAL FULFILLMENT OF THE AWARD OF THE DEGREE OF MASTER OF CLIMATE CHANGE ADAPTATION OF THE UNIVERSITY OF NAIROBI

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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.

Sampson, C., McEvoy, J., Oo, Z. M., Chit, A. M., Chan, A.N., Tonkyn, D., et al.
New elephant crisis in Asia—Early warning signs from Myanmar (2018)
PLoS ONE 13(3): e0194113. https://doi.org/10.1371/journal.pone.0194113

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In the southern Bago Yoma mountain range in Myanmar, Asian elephants are being killed at a disturbing rate. This emerging crisis was identified initially through a telemetry study when 7 of 19 of collared elephants were poached within a year of being fitted with a satellite-GPS collar. Subsequent follow up of ground teams confirmed the human caused death or disappearance of at least 19 elephants, including the seven collared individuals, within a 35 km2 area in less than two years. The carcasses of 40 additional elephants were found in areas located across south-central Myanmar once systematic surveys began by our team and collaborators. In addition to the extreme rate of loss, this study documents the targeting of elephants for their skin instead of the more common ivory, an increasing trend in Myanmar. Intensive research programs focused on other conservation problems identified this issue and are now encouraging local authorities to prioritize anti-poaching efforts and improve conservation policies within the country. Myanmar represents one of the last remaining countries in Asia with substantial wildlands suitable for elephants. Increasing rates of human-elephant conflict and poaching events in this country pose a dire threat to the global population.

Sarah McPherson, BBC Magazine.
Fighting for Elephants. (2015)
BBC Wildlife Magazine, January 2015 pp42-48

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Save the Elephants
2019 Annual Report (2019)
Save the Elephants

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Projects Updates from Save the Elephants

Save The Elephants
Living in harmony with elephants (2016)
Save The Elephants Education Program

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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.

Save The Elephants
Save The Elephants/Elephant Watch Scholarship Programme Booklet (2017)
Save The Elephants

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The Elephant Scholarship Programme is a primary branch of the Save The Elephants/Elephant Watch Safaris education program founded by Oria Douglas-Hamilton. We work closely with local primary schools in the Samburu Isiolo Conservation Area (SICA) to select intelligent students whose families are unable to afford school fees. Sponsoring these young people provides them with a rare and precious opportunity that will change their lives. Since its inception in 2001, the programme has benefitted over 150 students across Samburu, Isiolo and Marsabit counties. The programme goes the extra mile to build both intellect and character. This booklet is a brief chronicle of the different stages of some of our past and present scholarship students. Their unfaltering hearts, a thirst for education, humble backgrounds and unique experiences have transformed them into budding heroes and heroines of Save the Elephants and the community they live in.

Save The Elephants
2017 Annual Report (2017)
Save The Elephants

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Projects updates from Save The Elephants

Save the Elephants - The WILD Foundation - The Environment and Development Group
Initial Measures for Conservation of the Gourma Elephants, Mali. (2004)
Save the Elephants - The WILD Foundation - The Environment and Development Group ?Initial Measures for Conservation of the Gourma Elephants, Mali (2004) ?Save the Elephants-Interim Summary Progress Report.

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This report is summary in nature, and is submitted to keep the DNCN fully apprised of the progress of this project. The current Protocole d’Accord de Cooperation was signed in September 2003, and is in effect for two years. The parties involved have been:?• a consortium of The WILD Foundation (USA); Save the Elephants (STE- Kenya) and The Environment and Development Group (EDG - UK);?• the Direction Nationale de la Conservation de la Nature (Mali);?• with close collaboration by the Embassy of the United States in Bamako, Ambassador Vicki Huddleston.? For ease and brevity, this report will restate the overall mission and goals, then simply list the objectives of the 2004-05 field research and data compilation and briefly report on progress. Following that will be an outline summary of the 2005 work plan.

Save the Elephants - The WILD Foundation - The Environment and Development Group.
Mesures initiales pour la conservation des éléphants du Gourma, Mali. (2004)
Save the Elephants-Rapport d’activité sommaire intérimaire.

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Ce rapport est bref par nature et il est soumis afin que la DNCN reste pleinement informée des progrès de ce projet. Le Protocole d’Accord de Coopération actuel a été signé en septembre 2003 et est en vigueur pendant deux ans. Les parties concernées ont été :?• un consortium constitué de la Fondation WILD (E-U); de Save the Elephants (STE- Kenya) et du Environment and Development Group (EDG - GB) ;?• la Direction Nationale de la Conservation de la Nature (Mali) ;?• avec l’étroite collaboration de l’Ambassade des Etats-Unis à Bamako et notamment de l’Ambassadrice Vicki Huddleston.?Par souci de clarté et de brièveté, ce rapport réitère la mission d’ensemble et les buts du projet, puis dresse simplement la liste des objectifs des recherches de terrain et de collecte des données pour 2004-05 et donne un bref compte rendu des progrès réalisés. Un plan de travail sommaire sera ensuite présenté pour 2005.

Save the Elephants, ElephantVoices
Mara ecosystem connectivity Information on elephant population status and movements for spatial planning and conservation in Narok County (2016)
Save the Elephants and Elephant Voices.

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This document has been prepared at the invitation of the Narok County Government Department of Lands, Urban Development and Physical Planning and the County Assembly Committee on Natural Resources at a stakeholders forum on spatial plan development and resource mobilisation held in Narok on 16 - 17 October 2014. Kenya’s development blueprints - Vision 2030, the Constitution of Kenya (2010) and the Millennium Development Goals (MDG) - recognise the importance of sustainable use of natural resources, reduction of biodiversity loss, and maintenance of ecosystem processes. Vision 2030 identifies securing of wildlife dispersal areas and migratory routes/corridors and pathways as significant ingredients of sustainable eco-tourism. Biodiversity and ecosystem conservation contribute immensely to Kenya’s national economy with close to 80% of tourism earnings attributed to wildlife.

Sienne, J. M., Buchwald, R., Wittemyer, G.
Differentiation in mineral constituents in elephant selected versus unselected water and soil resources at Central African bais (forest clearings) (2014)
European Journal of Wildlife Research (2014) 60:377–382 DOI: 10.1007/s10344-013-0781-0

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Natural forest clearings (bais) in the Central African rain forest attract large numbers of mammals. Little is known about the factors influencing bai use by forest species, though geophagy and hydro-mineral resources are assumed to be important attractants. In the present study, clay and mineral concentrations in water and soil were examined at 15 bais. Water samples from elephant excavated pits showed significantly higher concentrations of most minerals sampled relative to surface waters. But mineral portfolios varied markedly between bais. Geophagy sites were less differentiated from control soil samples, leading to the interpretation that geophagy may not structure bai visitation. Monthly sampling of pit water at one bai suggested higher dry season mineral concentrations, which may relate to seasonal wildlife visitation patterns.

Soltis, J., King, L., Vollrath, F., Douglas-Hamilton, I.
Accelerometers and simple algorithms identify activity budgets and body orientation in African elephants Loxodonta africana (2016)
Endang Species Res 31: 1–12, 2016

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ABSTRACT: Accelerometers can be used to monitor animal behavior remotely, but validation is required for each species. Previously, we showed that accelerometer data in collars could be used to identify specific behaviors in African elephants Loxodonta africana, using complex analytical methods. Here, we show that simple methods can also be used to identify elephant activity levels and body orientation. Subjects were 6 African elephants: 3 at Disney’s Animal Kingdom®, Florida, USA, and 3 in Samburu-Laikipia, Kenya. Each elephant wore a collar containing a tri-axial ac - celerometer positioned on top of the neck. Simultaneous video recordings allowed validation of accelerometer data against observed behavior. The standard deviation of the total acceleration was shown to be a valid measure of dynamic acceleration, differentiating activity levels associated with resting, feeding, bathing, walking,

Soltis, J., Wilson, R.P., Douglas-Hamilton, I., Vollrath, F., King, L.E., Savage, A.
Accelerometers in collars identify behavioral states in captive African elephants Loxodonta africana. (2012)
Endang Species Res 18:255-263. doi: 10.3354/esr00452

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Accelerometers are motion-detection devices that, when attached to animals, are capable of detecting body orientation, overall activity levels, and specific behavior patterns. We deployed accelerometers in order to study the hypothesis that accelerometer output would allow us to distinguish between 4 behavior patterns in 3 adult female African elephants Loxodonta africana at Disney’s Animal Kingdom®, Florida, USA. Tri-axial accelerometer data loggers were attached to the tops of collars worn around the elephants’ necks. Behavior was documented on video while the accelerometer output was stored on the data logger. Feeding, bathing, and walking behaviors were recorded in all 3 subjects, while swaying behavior could be recorded in only 1 subject.

Taylor, L.A., Vollrath, F., Lambert, B., Lunn, D., Douglas-Hamilton, I., Wittemyer, G.
Movement reveals reproductive tactics in male elephants (2019)
J Anim Ecol. 2019;00:1–11 DOI: 10.1111/1365-2656.13035

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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.

Taylor, L.A., Wittemyer, G., Lambert, B., Douglas-Hamilton, I. and Vollrath, F.
Movement behaviour after birth demonstrates precocial abilities of African savannah elephant, Loxodonta africana, calves (2022)
Animal Behaviour, 187, pp.331-353.

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Living in groups has many benefits including predator protection and information sharing that can outweigh the associated costs. African savannah elephants live in a matriarchal fission–fusion social structure where groups merge or split depending on the relative costs of group living. However, elephants are also nonsynchronous breeders which may cause optimal movement patterns to differ between group members, particularly around parturition. Both the mother and the group must assess whether the benefits outweigh the costs of group cohesion. Understanding the impact of gestation, parturition and the presence of a newborn calf on the mother's movement patterns could provide key insights into the drivers of fission–fusion social dynamics. Here, we examined the relationship between reproductive state and movement using GPS tracking data collected during 23 parturitions from multiparous elephants (≥ second calving) and three from primiparous females. We analysed daily mean speed before, during and after parturition, and used a three-state hidden Markov model to determine whether parturition results in a distinctive change in speed. The results show that, although speed decreased on the estimated parturition date, parturition could only be detected in four of 23 multiparous parturition events when placed in the context of speed over a 2-month period. Furthermore, the speed on the day before and the day after parturition (i.e. first complete day of the calf's life) was not significantly different from the yearly mean.

Thouless, C., King, J., Omondi, P., Kahumbu, P., Douglas-Hamilton, I.
The status of Kenya’s elephants 1990–2002. (2002)
Kenya Wildlife Service

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The 1990s have been the first years since the 1960s that Kenya’s elephants have not substantially declined in numbers. Major savannah populations such as Tsavo, Laikipia–Samburu and Amboseli have increased significantly; others such as Mara and Meru have remained stable. Status of forest populations surveyed using dung counts is little known. Given the low confidence in such estimates, other indicators of population trends are employed. Forest area has declined, particularly in the major elephant ranges of Mt Kenya, the Aberdares and the Mau complex, as forests have been converted to farmland. There is no evidence that forest populations were affected by massive poaching of savannah-living elephants in the 1970s and 1980s. Density of most forest populations appear to be moderately high (more than 1 elephant per km2) and thus are unlikely to increase substantially.

Tiller, L., King, L., Lala, F., Pope, F., Thouless, C., Wall, J. and Douglas-Hamilton, I.
The outcome of an elephant translocation from Isiolo to Tsavo East National Park, Kenya (2022)
Pachyderm, 63, pp.91-98.

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ABSTRACT This management piece documents the outcome of an elephant translocation from Isiolo to Tsavo East National Park, Kenya in November 2021. The translocation aimed to reduce human-elephant conflict in the area and to prevent any retaliation toward the elephants. This was achieved by removing the ‘problem elephant group’, however the aim of resettling the elephants in Tsavo was not achieved, as the group fragmented and some swiftly moved far outside release site. Two elephants, which we were able to monitor through satellite collars, exhibited homing behaviour and both left Tsavo East National Park within 1-7 weeks of being released. If translocation continues to be the method of choice for problem elephants, there is a need for thorough planning and sound science to inform future operations, which should include collaring of each individual. Trained personnel and substantial budgeting for post release monitoring, and any potential conflict-reduction interventions, are therefore key management considerations for ensuring the health and wellbeing of translocated elephants in the future. In the long-term, focusing mitigation management on a larger number of habitual crop-raiders will have more impact and be a more effective approach for elephant managers. This could involve better spatial land-use planning, maintenance of corridors between protected areas, negative conditioning tactics and maintenance and upgrading of barriers.

Tiller, L.N., Humle, T., Amin, R., Deere, N.J., Lago, B.O., Leader-Williams, N., Sinoni, F.K., Sitati, N., Walpole, M., Smith, R.J.
Changing seasonal, temporal and spatial crop-raiding trends over 15 years in a human-elephant conflict hotspot (2021)
Biological Conservation Volume 254, February 2021, 108941 https://doi.org/10.1016/j.biocon.2020.108941

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Human-wildlife conflict is increasing due to rapid natural vegetation loss and fragmentation. We investigated seasonal, temporal and spatial trends of elephant crop-raiding in the Trans Mara, Kenya during 2014–2015 and compared our results with a previous study from 1999 to 2000. Our results show extensive changes in crop-raiding patterns. There was a 49% increase in incidents between 1999 -2000 and 2014–2015 but an 83% decline in the amount of damage per farm. Crop-raiding went from highly seasonal during 1999–2000 to year-round during 2014–2015, with crops being damaged at all growth stages. Additionally, we identified a new elephant group type involved in crop-raiding, comprising of mixed groups. Spatial patterns of crop-raiding also changed, with more incidents during 2014–2015 neighbouring the protected area, especially by bull groups. Crop-raiding intensity during 2014–15 increased with farmland area until a threshold of 0.4 km2 within a 1 km2 grid square, and farms within 1 km from the forest boundary, 2 km from village centres were most at risk of crop-raiding. In the last 20 years the Mara Ecosystem has been impacted by climate change, agricultural expansion and increased cattle grazing within protected areas. Elephants seem to have responded by crop-raiding closer to refuges, more frequently and throughout the year but cause less damage overall. While this means the direct economic impact has dropped, more farmers must spend more time protecting their fields, further reducing support for conservation in communities who currently receive few benefits from living with wildlife.

Tiller, L.N., King, L.E., Okita‐Ouma, B., Lala, F., Pope, F., Douglas‐Hamilton, I. and Thouless, C.R.
The behaviour and fate of translocated bull African savanna elephants (Loxodonta africana) into a novel environment (2022)
African Journal of Ecology, 60(4), pp.866-881.

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AbstractENTHIS LINK GOES TO A ENGLISH SECTIONFRTHIS LINK GOES TO A FRENCH SECTION Translocation of elephants is used to mitigate human-elephant conflict in Asia and Africa. However, few studies investigate how translocations affect the movements and social behaviour of individuals following their release, which may have important implications for whether translocated animals survive and succeed. Using GPS-tracking data, we explored movements of five translocated bull elephants (Loxodonta africana) moved to Tsavo, Kenya, and compared them with five resident bull elephants. Position data was collected hourly for 1 year (March 2018–March 2019), and analysed to investigate home range, displacement rates, problematic behaviour and group size. Of the five translocated elephants, three were illegally killed and one continued to break fences and raid crops. Only one elephant stayed away from human settlement. We found group size and composition to be significantly different, with translocated elephants observed in smaller groups with no female elephant interactions. All elephants showed variation in home ranges and displacement rates, but differences were not significant between resident and translocated elephant groups. For future translocations, we recommend careful consideration of elephant social systems, elephant age, timing, release site and proximity to human settlements that might create human-elephant conflict. This will improve chance for success of such high-stake and expensive translocations.

Tiller, L.N., Oniba, E., Opira, G., Brennan, E.J., King, L.E., Ndombi, V., Wanjala, D., Robertson, M.R.
“Smelly” Elephant Repellent: Assessing the Efficacy of a Novel Olfactory Approach to Mitigating Elephant Crop Raiding in Uganda and Kenya (2022)
Diversity 2022, 14, 509. https://doi.org/10.3390/d14070509

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Human–elephant conflict is increasing across many parts of Asia and Africa. Mitigating elephant crop raiding has become a major focus of conservation intervention, however, many existing methods for tackling this problem are expensive and difficult to execute. Thus, there is a need for more affordable, farm-based methods. Testing these methods is key to ensuring their effectiveness and feasibility. In this study, we tested a novel olfactory deterrent, the “smelly elephant repellent”, a foul-smelling organic liquid, on 40 farms in Uganda and Kenya. Our results show that the repellent was effective at deterring elephants from crop raiding. Over the study period, 82% of 309 elephant crop raids were deterred in Uganda. In Kenya, the repellent deterred 63% of 24 crop raiding incidents, and there was a significant effect of the repellent on test sites compared with control sites. The smelly repellent could be a helpful crop raiding mitigation tool for farmers, as this study showed it to be effective, relatively cheap, quick to produce from locally available ingredients, and communities have a positive attitude towards using it. Ongoing work is exploring the potential for a market-based approach to take this to scale in a financially sustainable way.

Tiller, L.N., Williams, H.F.
The elephant in the farm: long-term solutions are the key to coexistence (2021)
Anim. Conserv., 24: 733-734. https://doi.org/10.1111/acv.12741

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In this current state of exponential human population growth, natural spaces are being eroded more than ever before. Human activities have modified and transformed over half of the global land surface (Chapin et al., 2000), causing extensive habitat loss and fragmentation, and leading to a global decline in species. The rapid conversion of forest to agriculture puts farmers on the frontline of conflict with wildlife, and nowhere is this more pronounced than in the tropics, where development is rapidly catching up with the West. In the tropics communities are forced to survive alongside the megaherbivores that are predominantly under control or extinguished from much of the developed world. The most problematic animals for humans to live alongside are, arguably, elephants. Throughout much of elephant range in Asia and Africa, remaining elephant habitat is in the process of significant encroachment by humans, often with a front of edible crops that are highly preferred by such herbivores. These agricultural lands, on the edge of primary elephant habitat, can make for easily accessible resources for wildlife, and little is put in place to dissuade resident elephant herds from utilizing this resource. As a result, human-wildlife conflict is becoming a serious issue, and threatens the livelihoods of many of those living alongside megafauna, not to mention the threats to local elephant populations from retaliatory killings. Mitigating such situations often relies on translocating `problem animals’. In their paper, de la Torre et al. (2021) clearly show that such techniques are flawed. The overlap with elephants, in the peripheral agricultural landscapes, makes highly nutritious and abundant crops a readily available resource for elephants. Therefore, unless a constant stream of elephant translocations is a financially viable and sustainable solution, smarter methodologies need to be employed to create a model of coexistence where humans and elephants can live harmoniously together over the long term.

Troup, G., Doran, B., Au, J., King, L.E., Douglas-Hamilton, I., Heinsohn, R.
Movement tortuosity and speed reveal the trade-offs of crop raiding for African elephants (2020)
Animal Behaviour 168 (2020) 97e108 https://doi.org/10.1016/j.anbehav.2020.08.009

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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.

Troup, G., Heinsohn, R., King, L.E. and Edwards, K.L.
Exploring seasonal variation in the faecal glucocorticoid concentrations of African elephants (Loxodonta africana) living in a drought-prone, anthropogenic landscape (2022)
Wildlife Research - https://doi.org/10.1071/WR21003

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Context: The wide-ranging movement of African elephants (Loxodonta africana) is largely driven by the spatio-temporal distribution of water and forage, and often leads to their travelling outside of formally protected areas. With an increase in drier periods predicted across Africa due to climate change, it is critically important to understand how elephants physiologically respond to alterations in the availability and distribution of resources. Aims. We assessed variation in the adrenal activity of elephants living in Kenya’s Tsavo East National Park between wet and dry seasons, as well as among individuals found in Tsavo East National Park and privately protected Rukinga Wildlife Sanctuary (part of the Kasigau REDD+ wildlife corridor) in the dry season, when the area experiences an influx of elephants in search of alternative resources. Methods. We opportunistically collected fresh elephant faecal samples across the two seasons and locations for analysis of faecal glucocorticoid metabolite (fGCM, a proxy for stress) and nitrogen (Nf, an indirect measure of diet quality) concentrations. The Normalised Difference Vegetation Index (NDVI) was employed as an additional indicator of habitat quality. Key results. In Tsavo East N.P. Nf and NDVI were both significantly lower during the dry season, indicating poorer habitat quality compared with the wet season. Although elephant fGCM concentrations tended to be higher in the dry season than the wet, the differences were not significant. There was no difference between elephant fGCMs measured in Tsavo East N.P. and Rukinga W.S. during the dry season, nor in habitat quality between the two locations. Conclusions. Elephants living in Tsavo may be physiologically unaffected by (or adapt to) typical seasonal changes in habitat quality that could lead to nutritional stress; however, whether this is the case during extended periods of severe drought requires further investigation. Rukinga W.S. provides a safe haven of sufficient habitat quality for elephants searching for alternative resources during this period. Implications. Extended dry periods are likely to become increasingly common in semiarid savannahs, and implications for wildlife must be closely monitored. Privately protected land outside formally protected areas plays an important role in conservation efforts, which should be considered when making land management plans.

Tucker, M. A et al
Moving in the Anthropocene: Global reductions in terrestrial mammalian movements (2018)
Science. 2018 Jan 26;359(6374):466-469. doi: 10.1126/science.aam9712.

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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.

Turkalo, A. K., Wrege, P. H., Wittemyer, G.
Slow intrinsic growth rate in forest elephants indicates recovery from poaching will require decades (2016)
Journal of Applied Ecology © 2016 British Ecological Society

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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.

Turkalo, A.K., Wrege, P., Wittemyer, G.
Long-Term Monitoring of Dzanga Bai Forest Elephants: Forest Clearing Use Patterns. (2013)
PLoS One 8(12):e85154 DOI: 10.1371/journal.pone.0085154

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Individual identification of the relatively cryptic forest elephant (Loxodonta cyclotis) at forest clearings currently provides the highest quality monitoring data on this ecologically important but increasingly threatened species. Here we present baseline data from the first 20 years of an individually based study of this species, conducted at the Dzanga Clearing, Central African Republic. A total of 3,128 elephants were identified over the 20-year study (1,244 adults; 675 females, 569 males). It took approximately four years for the majority of elephants visiting the clearing to be identified, but new elephants entered the clearing every year of the study. The study population was relatively stable, varying from 1,668 to 1,864 individuals (including juveniles and infants), with increasingly fewer males than females over time.

Tyrrell, P., Amoke, I., Betjes, K., Broekhuis, F., Buitenwerf, R., Carroll, S., Hahn, N., Haywood, D., Klaassen, B., Løvschal, M. and Macdonald, D.
Landscape Dynamics (landDX) an open-access spatial-temporal database for the Kenya-Tanzania borderlands (2022)
Scientific Data, 9(1), pp.1-9 https://doi.org/10.1038/s41597-021-01100-9

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Abstract The savannas of the Kenya-Tanzania borderland cover >100,000 km2 and is one of the most important regions globally for biodiversity conservation, particularly large mammals. The region also supports >1 million pastoralists and their livestock. In these systems, resources for both large mammals and pastoralists are highly variable in space and time and thus require connected landscapes. However, ongoing fragmentation of (semi-)natural vegetation by smallholder fencing and expansion of agriculture threatens this social-ecological system. Spatial data on fences and agricultural expansion are localized and dispersed among data owners and databases. Here, we synthesized data from several research groups and conservation NGOs and present the first release of the Landscape Dynamics (landDX) spatial-temporal database, covering ~30,000 km2 of southern Kenya. The data includes 31,000 livestock enclosures, nearly 40,000 kilometres of fencing, and 1,500 km2 of agricultural land. We provide caveats and interpretation of the different methodologies used. These data are useful to answer fundamental ecological questions, to quantify the rate of change of ecosystem function and wildlife populations, for conservation and livestock management, and for local and governmental spatial planning.

Uno, K. T., Fisher, D. C., Wittemyer, G., Douglas-Hamilton, I., Carpenter, N., Omondi, P., Cerling, T. E.
Forward and inverse methods for extracting climate and diet information from stable isotope profiles in proboscidean molars (2020)
Quaternary International, Volume 557, 2020, Pages 92-109, ISSN 1040-6182 https://doi.org/10.1016/j.quaint.2020.06.030

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Abstract: Intratooth stable isotope profiles in enamel provide time series of dietary and environmental information that if correctly interpreted, serve as archives of seasonal variability in past environments. A major challenge in interpreting these profiles arises from time averaging imparted by enamel mineralization and developmental geometry, whereby the primary (δ13C or δ18O) input signal is attenuated and shifted, which can potentially lead to incorrect interpretations of the magnitude or frequency of seasonal variability. Several forward and inverse models have been developed to reconstruct the primary input signal from intratooth profiles in continuously growing teeth. Here the models developed by Passey and Cerling (2002) and Passey et al. (2005) are extended to molars of Elephantinae, which grow over a long but finite interval of time. Proboscidean molars are particularly attractive for intratooth profiles because they may contain a decade or more of information and they are often well preserved in the fossil record because of their thick enamel and large size. Forward model parameters are established using histological analysis of molar thin sections of extant African elephants (Loxodonta africana) and a mammoth (Mammuthus columbi) and by micro-CT analysis of L. africana molar plates. The density of immature enamel is about 65% of the final density of mature enamel. The appositional length varies from approximately 35 to 55 mm, and the maturation length is about 70 mm. Histological methods are used to determine crown formation time (CFT) in elephant and mammoth molar plates. CFT for the elephant and mammoth molar plates studied in thin section are about 5–6 years and 11 years, which translate to mean growth rates of about 21 mm/year and 16 mm/year, respectively. Coeval molar and tusk profiles from a zoo elephant are compared. The tusk isotope profile serves as a proxy for the primary input signal, and thus provides an opportunity to evaluate the forward and inverse models. The results from the zoo elephant profiles demonstrate that the inverse model accurately reconstructs the amplitude and overall structure of the primary input signal. Inverse model results of mammoth molar profiles show double the range of δ13C in measured enamel profiles. Inversion model results illustrate that improved reconstruction of the primary input signal can lead to more accurate interpretations of the seasonal variability of diet and body water and by extension, vegetation and precipitation in past environments.

Uno, K.T., Quade, J., Fisher, D.C., Wittemyer, G., Douglas-Hamilton, I., Andanje, S., Omondi, P., Litoroh, M., Cerling, T.E.
Bomb-curve radiocarbon measurement of recent biologic tissues and applications to wildlife forensics and stable isotope (paleo) ecology. (2013)
PNAS 2013 ; published ahead of print July 1, 2013, doi:10.1073/pnas.1302226110

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Above-ground thermonuclear weapons testing from 1952 through 1962 nearly doubled the concentration of radiocarbon (14C) in the atmosphere. As a result, organic material formed during or after this period may be radiocarbon-dated using the abrupt rise and steady fall of the atmospheric 14C concentration known as the bomb-curve. We test the accuracy of accelerator mass spectrometry radiocarbon dating of 29 herbivore and plant tissues collected on known dates between 1905 and 2008 in East Africa. Herbivore samples include teeth, tusks, soft tissue, hair, and horn. Tissues formed after 1955 are dated to within 0.3–1.3 y of formation, depending on the tissue type, whereas tissues older than ca. 1955 have high age uncertainties (>17 y) due to the Suess effect. 14C dating of tissues has applications to stable isotope (paleo) ecology and wildlife forensics.

van de Water, A., King, L. E., Arkajak, R., Arkajak, J., van Doormaal, N., Ceccarelli, V., Sluiter, L., Doornwaard, S. M., Praet, V., Owen, D., Matteson, K.
Beehive fences as a sustainable local solution to human‐elephant conflict in Thailand (2020)
Conservation Science and Practice. 2020;e260 https://doi.org/10.1111/csp2.260

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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.

Vigne, L., Martin, E
Myanmar’s Growing Illegal Ivory Trade With China (2018)
Save The Elephants

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Vigne, L., Martin, E.
VIETNAM’S ILLEGAL IVORY TRADE THREATENS AFRICA’S ELEPHANTS. (2016)
Save the Elephants

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Art historians are uncertain when the Vietnamese first started to use ivory in any significant manner within the country. Certain tribal ethnic groups had used a small amount of ivory from the elephants living in the forests around them, some of which they captured and domesticated. The Mnong tribe, for example, are well known historically for wearing ivory earplugs, still seen for sale in antique shops in the main cities, but silver jewellery now is the traditional symbol of wealth in many hill tribes in the region, such as silver necklaces and headdress ornaments. The use of ivory is minor, in comparison. Raw tusks have passed through Vietnam to China for hundreds of years. In the early 15th century King Le Loi of Vietnam sent four pieces of tusk to China as part of a peace pact (Nguyen-Long 2013). The tradition in Vietnam is long, dating back over 1,000 years of Vietnamese artisans

Vigne, L., Martin, E.
China faces a conservation challenge: The Expanding Elephant and Mammoth Ivory Trade in Beijing and Shanghai (2014)
Save the Elephants & Aspinall Foundation

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The Chinese ivory industry has been expanding tremendously, especially in the last five years. Illegal ivory imports are the largest in the world by weight and have been soaring; the wholesale price of raw tusks sold for carving has tripled since 2010; the number of official ivory carving factories has risen several-fold; the number of carvers is way up, the number of legal (licensed) retail outlets for elephant ivory has increased significantly as has the number of illegal (unlicensed) outlets selling elephant ivory; there has been a boom in customers for elephant ivory, both legal and illegal, both raw and worked, especially by collectors and investors in China; retail prices for ivory items have skyrocketed; and some businesspeople are optimistic about future economic benefits of trading in ivory.

Vigne, L., Martin, E.
Decline in the legal ivory trade in China in anticipation of a ban (2017)
Save The Elephants

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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

Vigne, L., Martin, E.
THE IVORY TRADE OF LAOS: NOW THE FASTEST GROWING IN THE WORLD (2017)
SAVE THE ELEPHANTS

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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.

Vigne, L., Martin, E.
BUÔN LAU NGÀ VOI O VIET NAM DANG DE DOA TOI VOI CHÂU PHI. (2016)
Save the Elephants

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Theo m?t báo cáo m?i phát hành ngày hôm nay c?a T? ch?c Save the Elephants, Vi?t Nam hi?n là m?t trong nh?ng th? tr??ng kinh doanh ngà voi trái phép l?n nh?t trên th? gi?i. Qua th?ng kê, s? l??ng s?n ph?m ???c buôn bán ?ã t?ng h?n sáu l?n t? n?m 2008 ??n n?m 2015. Ngoài Vi?t Nam, ch?a có qu?c gia nào ???c ghi nh?n có các ho?t ??ng trong c? hai l?nh v?c: nh?p kh?u trái phép các lo?i ngà voi d?ng thô và xu?t kh?u trái phép các thành ph?m t? ngà voi. T?ng c?ng, có t?i 242 ?i?m tiêu th?/ ??i ly?, bày bán 16.099 s?n ph?m có ngu?n g?c t? ngà voi cho các ??i t??ng khách hàng bán l?, ???c phát hi?n t?i TP. H? Chí Minh (TP. HCM), TP. Buôn Ma Thu?t, Hà N?i và các khu v?c làng xã lân c?n. Theo m?t báo cáo công b? n?m 2008, s? l??ng s?n ph?m ngà voi bày bán ch? ??t m?c 2.444 s?n ph?m.

Vollrath, F.
Trunks,tracks and spiders’ webs. (2007)
Oxford Today . Hilary Issue

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In Roman times, elephants roamed widely over most of Africa and Asia, probably linking up somewhere around Mesopotamia. Indeed, not very much earlier, the elephant tribe consisted of hundreds of species ranging even further afield, covering most of Europe and America as well – with mammoths of up to five metres in the icy north and one-metre dwarf forms on Mediterranean (and Californian) islands. Everywhere, the large beasts shaped their environment by pushing over trees and denuding forests, thus creating open grasslands. Those halcyon days are long past, and all but two species are now extinct. Times have become rough for the survivors: over the past 100 years, the Asian Elephas maximus and the African Loxodonta africana have had to yield to human expansion and retreat into a few small pocket-size remnants of their natural ranges.

Vollrath, F., Douglas-Hamilton, I.
African bees to control African elephants. (2002)
Naturwissenschaften 89, 508-511

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Numbers of elephants have declined in Africa and Asia over the past 30 years while numbers of humans have increased, both substantially. Friction between these two keystone species is reaching levels which are worryingly high from an ecological as well as a political viewpoint. Ways and means must be found to keep the two apart, at least in areas sensitive to each species' survival. ??The aggressive African bee might be one such method. Here we demonstrate that African bees deter elephants from damaging the vegetation and trees which house their hives.

Wall, J., Douglas-Hamilton, I., Vollrath, F.
Elephants Avoid Costly Mountaineering. (2006)
Current Biology July 2006 Vol. 16, Issue 14. R527-9

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Understanding the behavioural decisions underlying animal movements is a major challenge. Here we report evidence for the importance of the abiotic terrain feature ‘gradient’ in guiding the movements of African savannah elephants (Loxodonta africana). Global Positioning System (GPS) tracking data overlaid onto digital elevation and surface gradient models show that elephants tend to avoid steep slopes. Energy calculations suggest that even minor hills are considerable energy barriers for heavy animals. Elephants are keystone animals in Africa and Asia, and effective conservation planning strategies must integrate a thorough knowledge of the range use and spatial requirements of these magnificent animals.

Wall, J., Wittemyer, G., Klinkenberg, B., Douglas-Hamilton, I.
Novel opportunities for wildlife conservation and research with real-time monitoring. (2014)
Ecological Applications Volume 24, Issue 4 (June 2014) pp. 593-601 doi: http://dx.doi.org/10.1890/13-1971.1

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The expansion of global communication networks and advances in animal tracking technology make possible the real-time telemetry of positional data as recorded by animal-attached tracking units. When combined with continuous, algorithm-based analytical capability, unique opportunities emerge for applied ecological monitoring and wildlife conservation. We present here four broad approaches for algorithmic wildlife monitoring in real time—proximity, geofencing, movement rate, and immobility—designed to examine aspects of wildlife spatial activity and behavior not possible with conventional tracking systems. Application of these four routines to the real-time monitoring of 94 African elephants was made. We also provide details of our cloud-based monitoring system including infrastructure, data collection, and customized software for continuous tracking data analysis.

Wall, J., Wittemyer, G., Klinkenberg, B., LeMay, V., and Douglas-Hamilton, I.
Characterizing Properties and Drivers of Long Distance Movements by Elephants (Loxodonta africana) in the Gourma, Mali. (2013)
Biological Conservation 157: 60-68. Online 2012.  doi.org/10.1016/j.biocon.2012.07.019

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Wide-ranging, landscape-level movements by terrestrial herbivores are increasingly threatened globally. Understanding the ecology of spatio-temporal movement patterns is critical for conservation of wide-ranging terrestrial species and the ecosystems on which they rely. The range of the Gourma elephant population inhabiting the Sahelian eco-region near Tombouctou (Timbuktu), Mali encompasses the largest areal extent in this species (29% greater than range sizes reported in other populations). Over the course of a year, the Gourma elephants (Loxodonta africana) move in a coordinated north–south movement pattern that is relatively unique for the species.

Wall, J., Wittemyer, G., Klinkenberg, B., LeMay, V., Blake, S., Strindberg, S., Henley, H., Vollrath, F., Maisels, F., Ferwerda, J., Douglas-Hamilton, I.
Human footprint and protected areas shape elephant range across Africa (2021)
Current Biology 31, 1–9, June 7, 2021 DOI:https://doi.org/10.1016/j.cub.2021.03.042

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Summary Over the last two millennia, and at an accelerating pace, the African elephant (Loxodonta spp. Lin.) has been threatened by human activities across its range.1, 2, 3, 4, 5, 6, 7 We investigate the correlates of elephant home range sizes across diverse biomes. Annual and 16-day elliptical time density home ranges8 were calculated by using GPS tracking data collected from 229 African savannah and forest elephants (L. africana and L. cyclotis, respectively) between 1998 and 2013 at 19 sites representing bushveld, savannah, Sahel, and forest biomes. Our analysis considered the relationship between home range area and sex, species, vegetation productivity, tree cover, surface temperature, rainfall, water, slope, aggregate human influence, and protected area use. Irrespective of these environmental conditions, long-term annual ranges were overwhelmingly affected by human influence and protected area use. Only over shorter, 16-day periods did environmental factors, particularly water availability and vegetation productivity, become important in explaining space use. Our work highlights the degree to which the human footprint and existing protected areas now constrain the distribution of the world’s largest terrestrial mammal.9,10 A habitat suitability model, created by evaluating every square kilometer of Africa, predicts that 18,169,219 km2 would be suitable as elephant habitat—62% of the continent. The current elephant distribution covers just 17% of this potential range of which 57.4% falls outside protected areas. To stem the continued extirpation and to secure the elephants’ future, effective and expanded protected areas and improved capacity for coexistence across unprotected range are essential.

Wall, J., Wittemyer, G., LeMay, V., Douglas-Hamilton, I., Klinkenberg, B.
Elliptical Time-Density model to estimate wildlife utilization distributions. (2014)
Methods in Ecology and Evolution http://onlinelibrary.wiley.com/doi/10.1111/2041-210X.12218/abstract

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We present a new animal space-use model (Elliptical Time-Density - ETD) that uses discrete-time tracking data collected in wildlife movement studies. The ETD model provides a trajectory-based, non-parametric approach to estimate the utilization distribution (UD) of an animal, using model parameters derived directly from the movement behavior of the species. The model builds on the theory of ‘time-geography’ whereby elliptical constraining regions are established between temporally-adjacent recorded locations. Using a Weibull speed distribution fitted for an animal's movement data, a time-density value (i.e., time per unit landscape) is determined from the expectation of all elliptical regions equal to, or greater-than, the minimum bounding ellipse for a given landscape point.

Wasser, S., et al
Elephants, Ivory, .and Trade (2010)
Science Vol 327

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Wasser, S., Poole, J., Lee, P., Lindsay, K., Dobson, A., Hart, J., Douglas-Hamilton, I., Wittemyer, G., Granli, P., Morgan, Bethan., Gunn, J., Alberts, S., Beyers, R., Chiyo, P., Croze, H., Estes, R., Gobush, K., Joram, P., Kikoti. A., Kingdon, J., King, L., Macdonald, D., Moss, C., Mutayoba, B., Njumbi, S., Omondi, P., Nowak, K Tanzania and Zambia are petitioning the Convention on International Trade in Endangered Species (CITES) to “downlist” the conservation status of their elephants to allow sale of stockpiled ivory. But just 2 years after CITES placed a 9-year moratorium on future ivory sales, elephant poaching is on the rise. The petitioning countries are major sources and conduits of Africa’s illegal ivory. The petitions highlight the controversy surrounding ivory trade and broader issues underlying CITES trade decisions.

Weideman, H., Stewart, C., Parham, J., Holmberg, J., Flynn, K., Calambokidis, J., Paul, D, B., Bedetti, A., Henley, M., Lepirei, J., Pope, F.
Extracting identifying contours for African elephants and humpback whales using a learned appearance model (2020)
2020 IEEE Winter Conference on Applications of Computer Vision (WACV)

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Abstract This paper addresses the problem of identifying individual animals in images based on extracting and matching contours, focusing in particular on the trailing edges of humpback whale flukes and the outline of the ears of African savanna elephants. A coarse-grained FCNN is learned to isolate the contour in an image, and a fine-grained FCNN is learned to provide more precise boundary information. The latter is trained by generating synthetic boundaries from coarse, easily-extracted training data, avoiding tedious manual effort. An A* algorithm extracts the final contour, which is converted to set of digital curvature descriptors and matched against a database of descriptors using local-naive Bayes nearest neighbors. We show that using the learned fine-grained FCNN produces more accurate contours than using image gradients for fine localization, especially for elephant ears where the boundaries are primarily texture. Matching using contours extracted using the fine-grained FCNN improves top-1 accuracy from 80% to 85% for flukes and 78% to 84% for ears.

Weinmann, S.
Impacts of Elephant Crop-Raiding on Subsistence Farmers and Approaches to Reduce Human- Elephant Farming Conflict in Sagalla, Kenya (2018)
Thesis presented in partial fulfillment of the requirements for the degree of Master of Science In Resource Conservation, International Conservation and Development The University of Montana, Missoula, MT

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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.

Wilkie, R.D., and D-H, I.
High-resolution tracking technology reveals distinct patterns in nocturnal crop raiding behaviour of an African elephant (Loxodonta africana) in Amboseli, Kenya (2018)
Pachyderm No. 59 July 2017–June 2018

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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.

Williams, H., Leneuyia, K., Mwalavo, B., Serem, G., Sempeyo, V., Nyakundi, B., Tiller, L., Pope, F. and King, L.
Part II: Using cinema, theatre and a virtual toolbox to address the conflict between humans and elephants (2022)
Pachyderm, 63, pp.158-164. July - Sept 2022

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ABSTRACT The value of art in Science is undisputed. From DaVinci’s drawings came a plethora of inventions, most notable is arguably his “aerial screw” which is highly suggestive of the helicopter we know today (Da Vinci, 1894), and the night skies of Charles Mezzier who, through his drawings documented countless formerly undocumented celestial bodies (Messier, 1781). In these and many other cases great art has undoubtedly gone on to aid science. Within the field of conservation the benefits of art are more nuanced. Recently art in its modern forms has been proven to be impactful in terms of attitudes to nature with multiple nature documentaries being empirically proven to have a range of impacts (Silk et al., 2021, Jones et al., 2019), from knowledge gains not translating into reduction in plastic usage following the viewing of Blue Planet II (Dunn et al., 2020) to Blackfish causing a decrease in the market value of Seaworld (Boissat et al., 2021). Animal imagery including photos and drawings as well as movies have also been shown to generally improve attitudes to nature (Thomas-Walters et al., 2020). But does art really have a place in modern day conservation where more species than ever before are on the brink of extinction due to human activities (Barnosky et al., 2011)?

Wittemyer, G.
Effects of Economic Downturns on Mortality of Wild African Elephants. (2011)
Conservation Biology. doi:10.1111/j.1523-1739.2011.01713.x

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Declines in economic activity and associated changes in human livelihood strategies can increase threats of species overexploitation. This is exemplified by the effects of economic crises, which often drive intensification of subsistence poaching and greater reliance on natural resources. Whereas development theory links natural resource use to social-economic conditions, few empirical studies of the effect of economic downturns on wild animal species have been conducted. I assessed the relations between African elephant (Loxodonta africana) mortality and human-caused wounds in Samburu, Kenya and (1) livestock and maize prices (measures of local economic conditions), (2) change in national and regional gross domestic product (GDP) (measures of macroeconomic conditions), and (3) the normalized difference vegetation index (NDVI) (a correlate of primary productivity).

Wittemyer, G.
The Socio-Ecology of the African Elephant (Loxodonta africana) (2005)
PhD dissertation. GRADUATE DIVISION of the UNIVERSITY OF CALIFORNIA, BERKELEY

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A variety of challenges face the conservation of African elephants, stemming from the illegal poaching for ivory to habitat loss resulting in range restriction. Solutions to these challenges require information on the factors affecting population structure, movement and reproduction in this species. In this dissertation, I investigate the relationship between ecological variation and population processes in the wild elephant population inhabiting the Samburu and Buffalo Springs National Reserves in northern Kenya. Both empirical analyses and theoretical approaches are presented, motivated by fundamental questions regarding factors influencing population structure and by applied objectives concerning the management issues facing this species. In addition, this work presents novel analytical techniques for defining and understanding population structure.

Wittemyer, G.
The elephant population of the Samburu and Buffalo Springs National Reserves, Kenya.  (2001)
Afr. J. Ecol. 39, 357-365

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A 21-month individual identification project on the Samburu and Buffalo Springs National Reserves’ elephant population was conducted between November 1997 and July 1999. The free ranging population, of at least 767 elephants, which relied heavily on areas outside the reserves, was individually identified. The numbers of elephants observed per day fluctuated but were greater during dry periods then wet. However, the sizes of aggregations were greater during wet periods. Preliminary investigation suggested that the population could be divided into two groups, which were designated resident and non-resident family units. The groups comprised approximately equal numbers of cows and calves, but temporally had di¡erent reserve use patterns and calving peaks. The daily numbers of males and musth males were correlated with numbers of females.

Wittemyer, G.
The Socio-Ecology of the African Elephant (Loxodonta africana) (2005)
A dissertation submitted in partial satisfaction of the requirements for the degree of Doctor of Philosophy in Environmental Science, Policy and Management in the Graduate Division of the University of California, Berkeley

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A variety of challenges face the conservation of African elephants, stemming from the illegal poaching for ivory to habitat loss resulting in range restriction. Solutions to these challenges require information on the factors affecting population structure, movement and reproduction in this species. In this dissertation, I investigate the relationship between ecological variation and population processes in the wild elephant population inhabiting the Samburu and Buffalo Springs National Reserves in northern Kenya. Both empirical analyses and theoretical approaches are presented, motivated by fundamental questions regarding factors influencing population structure and by applied objectives concerning the management issues facing this species.

Wittemyer, G.
Stemming Wildlife Poaching. (2015)
Testimony for the Senate Foreign Relations Subcommittee on Africa and Global Health Policy. US Government Printing Office

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Testimony for the Senate Foreign Relations Subcommittee on Africa and Global Health Policy Hearing on “Stemming Wildlife Poaching” - Thursday, July 16, 2015 Mr. Chairman, Mr. Ranking Member, and members of the Committee, I want to thank you for the opportunity to submit testimony for the record of this hearing. I am honored to appear before your committee. My name is George Wittemyer—I am a professor in the Department of Fish, Wildlife and Conservation Biology at Colorado State University and the Chairman of the Scientific Board of the Kenya based organization Save the Elephants. I have worked on elephant conservation issues in Africa for the past 19 years and have been a member of the IUCN African Elephant Specialist Group for the past 8 years. In addition, I serve as a technical advisor on elephants to the Kenya Wildlife Service.

Wittemyer, G., Cerling, T.E., Douglas-Hamilton, I.
Establishing chronologies from isotopic profiles in serially collected animal tissues: An example using tail hairs from African elephants. (2009)
Chemical Geology 267 (2009) 3–11

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While the use of stable isotopes in wildlife ecological research is growing rapidly, development of methods to establish time-specific isotope data from continuously growing animal tissues are lacking. Using serially collected tail hairs from wild African elephants (Loxodonta africana), we develop and compare four techniques to collate temporal isotope chronologies from metabolically inert tissues for which formation/growth overlapped in time. The influence of variation in within hair growth rates and other sources of error in the presented techniques are explored and found to be inconsequential relative to the 5-day tissue sampling interval. Using a floating point regression approach, we find a high degree of correlation between independently derived isotope profiles from the same and different individuals in the study ecosystem.

Wittemyer, G., Daballen, D., Douglas-Hamilton, I
Differential influence of human impacts on age-specific demography underpins trends in an African elephant population (2021)
Ecosphere 12(8):e03720. 10.1002/ecs2.3720 https://doi.org/10.1002/ecs2.3720 August 2021

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Abstract Diagnosing age-specific influences on demographic trends and their drivers in at-risk wildlife species can support the development of targeted conservation interventions. Such information also underpins understanding of life history. Here, we assess age-specific demography in wild African elephants, a species whose life history is marked by long life and extreme parental investment. During the 20-yr study, survival and its variation were similar between adults and juveniles in contrast to relationships found among many large-bodied mammals. Prospective analysis on age-specific Leslie matrices for females demonstrated survival is more influential than fecundity on λ, with sensitivity of both decreasing with age. Results aggregated by stage classes indicate young adults (9–18 yr) demonstrated the highest elasticity, followed by preparous juveniles (3–8 yr). Mature adults (36+ yr) had the lowest aggregate elasticity value. Retrospective analysis parameterized by data from the early and latter periods of the study, characterized by low then high human impact (faster and slower growth, respectively), demonstrated fecundity (particularly for adults; 19–35 yr) explained the greatest variation in λ observed during the period of low human impact, while survival (particularly juvenile and adult) was more influential during the high human impact period. The oldest females (mature adult stage) weakly influenced population growth despite demonstrating the highest fecundity and their behavioral importance in elephant society. Multiple regression models on survival showed the negative effects of human impacts and population size were the strongest correlates across sexes and ages. Annual rainfall, our metric for environmental conditions, was weakly informative. The presence of dependent young was positively correlated with survival for breeding females, suggesting condition-based mortality filtering during pregnancy. Notwithstanding the stabilizing effect of high juvenile survival on elephant population growth, demographic processes in elephants were similar to those shaping life history in other large herbivores. Implications of the study results with respect to the conservation of elephants and analysis of demographic impact of poaching are discussed, along with the study's relevance to theories regarding the evolution of life history and parental care.

Wittemyer, G., Daballen, D., Douglas-Hamilton, I.
Comparative Demography of an At-Risk African Elephant Population. (2013)
PLoS ONE 8(1): e53726. doi:10.1371/journal.pone.0053726

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Knowledge of population processes across various ecological and management settings offers important insights for species conservation and life history. In regard to its ecological role, charisma and threats from human impacts, African elephants are of high conservation concern and, as a result, are the focus of numerous studies across various contexts. Here, demographic data from an individually based study of 934 African elephants in Samburu, Kenya were summarized, providing detailed inspection of the population processes experienced by the population over a fourteen year period (including the repercussions of recent increases in illegal killing). These data were compared with those from populations inhabiting a spectrum of xeric to mesic ecosystems with variable human impacts.

Wittemyer, G., Daballen, D., Douglas-Hamilton, I.
Rising ivory prices threaten elephants. (2011)
Nature 476, 282-283 (18 August 2011). doi:10.1038/476282c

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Kenya’s premier Samburu elephant population is the focus of a distressing surge in ivory poaching, coincident with an increase in illegal trading of ivory. This ivory is mainly destined for China (see go.nature.com/czac3x). Effective protection of elephants depends partly on more conservation investment, but mainly on stemming the demand for ivory and eliminating black-market trade — actions that mandate leadership from and cooperation with China. The Samburu elephants are one of the world’s best-studied populations. Intensive monitoring has revealed that more have been poached in the past 2.5 years than in the previous 11 years. The highest poaching rates ever were recorded in the first 5 months of this year.

Wittemyer, G., Daballen, D., Rasmussen, H., Kahindi O., Douglas-Hamilton, I.
Demographic status of elephants in the Samburu and Buffalo Springs National Reserves, Kenya. (2005)
African Journal of Ecology, Afr. J. Ecol., 43, 44–47

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Individual based demographic records of the elephants utilizing Samburu and Buffalo Springs National Reserves were collected from 1998 through 2003 and indicate that this elephant population was increasing at an average rate of 4.6% per year. Although the majority of carcasses were not found, known sources of mortality include disease,injury, and predation by lions and humans. Poaching did occur during the study period, however the population is increasing and thus our findings indicate ivory poaching has limited impact on the demographic status of these elephants. This population is part of the Samburu/Laikipia MIKE Site and thus its status is relevant to CITES legislation.

Wittemyer, G., Douglas-Hamilton, I., Getz, W.
The socioecology of elephants: analysis of the processes creating multitiered social structures. (2005)
Animal Behaviour 69, 1357-1371. 2005

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In this paper, we investigate the formation and function of the multilevelled, fission-fusion social structure in a free-ranging African elephant, Loxodonta africana, population. We quantitatively identified the existence of four social tiers by using cluster analysis on individual association data. We assessed the effects of season and study period on social structuring and levels of cohesion within and among social units. ??We found that second-tier units, potentially the equivalent of the 'family', were stable across seasonal periods but the number of units increased as the study progressed and the population grew. It appears that these units were sufficiently small not to be influenced by ecologically related factors, such as resource competition, that might otherwise lead to them splitting.

Wittemyer, G., Ganswindt, A., Hodges, K.
The impact of ecological variability on the reproductive endocrinology of wild female African elephants. (2007)
Elsevier. Hormones and Behavior 51 (2007) 346–354

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Non-invasive endocrine methods enable investigation of the relationship between ecological variation and ovarian activity and how this impacts on demographic processes. The underlying physiological factors driving high variation in inter-calving intervals among multi-parous African elephants offer an interesting system for such an investigation. This study investigates the relationship between Normalized Differential Vegetation Index (NDVI), an ecosystem surrogate measure of primary productivity, and fecal progestin concentrations among wild female elephants. Matched fecal samples and behavioral data on reproductive activity were collected from 37 focal individuals during the two-year study.

Wittemyer, G., Getz, W. M.
Hierarchical dominance structure and social organization in African elephants, Loxodonta africana. (2007)
Elsevier. The Association for the Study of Animal Behaviour

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According to the socioecological framework, transitivity (or linearity) in dominance relationships is related to competition over critical resources. When a population is structured into groups, the intensity of between versus within-group competition influences the form and function of its social organization. Few studies have compared the type and relative intensity of competition at these two levels. African elephants have well-structured social relations, providing an exemplary system for such a study. We report on dominance hierarchies among free-ranging elephants and evaluate the factors that drive their socioecological structure to lie in a region of the three-dimensional nepotism/despotism/tolerance space rarely observed among social species;

Wittemyer, G., Getz, W.M.
A likely ranking interpolation for resolving dominance orders in systems with unknown relationships. (2006)
Koninklijke Brill NV, Leiden, 2006. Behaviour 143, 909-930

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In many animal systems agonistic interactions may be rare or not overt, particularly where?such interactions are costly or of high risk as is common for large mammals. We present a?technique developed specifically for resolving an optimized dominance order of individuals?in systems with transitive (i.e. linear) dominance relationships, but where not all relationships?are known. Our method augments the widely used I&SI method (de Vries, 1998) with?an interpolation function for resolving the relative ranks of individuals with unknown relationships.?Our method offers several advantages over other dominance methods by enabling?the incorporation of any proportion of unknown relationships, resolving a unique solution to?any dominance matrix, and calculating cardinal dominance strengths for each individual. As?such, this method enables novel insight into difficult to study behavioural systems.

Wittemyer, G., Keating, L. M., Vollrath, F., Douglas-Hamilton, I.
Graph theory illustrates spatial and temporal features that structure elephant rest locations and reflect risk perception (2017)
Ecography, 40: 598–605. doi:10.1111/ecog.02379

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Understanding the spatial structuring of animal behaviors and how they link landscapes can be critical for conservation management. This emerging field has been greatly facilitated by technologically advanced acquisition and analysis of data on animal movements. The framework of graph theory, which directly quantifies network connectivity properties, provides a useful addition to this tool set. Using a novel application of graph theory, we investigate the structure and patterning of African elephant Loxodonta africana rest sites, a potentially critical feature structuring spatial properties of animal populations. Elephants in the study rested intermittently and for short durations (1–3 rests d–1, lasting 3–5 h total). They switched circadian rest patterns according to landscape attributes, resting more during the day and further from permanent water in areas with high human density outside protected areas. Within protected areas and during the dry season, elephants showed clustering and sequential use of rest nodes (repeated motifs). Repeated use of specific rest nodes (self-looping) was more frequent than expected if rest nodes were chosen at random, particularly when outside protected areas further from water, indicating the importance of preferred rest sites. Our results suggest that elephants adjust resting behavior when in human-dominated areas, using preferred resting sites presumably in locations that reduce the risk of interactions. This study demonstrates how graph theory may be used practically to gain novel insight into behaviours, such as resting, that are discrete in time and space. Furthermore, analysis of the spatial and network properties of rest sites, given an individual's susceptibility when engaged in rest behavior, allowed characterization of spatio-temporal risk perception, providing a powerful behavioral based means to quantify the landscape of fear.

Wittemyer, G., Northrup, J.M., Blanc, J., Douglas-Hamilton, I., Omondi, P., Burnham, K.P.
Illegal killing for ivory drives global decline in African elephants (2014)
Wittemyer, G., Northrup, J.M., Blanc, J., Douglas-Hamilton, I., Omondi, P., Burnham, K.P.

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Illegal wildlife trade has reached alarming levels globally, extirpating populations of commercially valuable species. As a driver of biodiversity loss, quantifying illegal harvest is essential for conservation and sociopolitical affairs but notoriously difficult. Here we combine field-based carcass monitoring with fine-scale demographic data from an intensively studied wild African elephant population in Samburu, Kenya, to partition mortality into natural and illegal causes. We then expand our analytical framework to model illegal killing rates and population trends of elephants at regional and continental scales using carcass data collected by a Convention on International Trade in Endangered Species program.

Wittemyer, G., Okello, J.B.A., Rasmussen, H.B., Arctander, P., Nyakaana, S., Douglas-Hamilton, I., Siegismund, H.
Where sociality and relatedness diverge: the genetic basis for hierarchical social structure in African elephants. (2009)
Proceedings of the Royal Society: Biological Sciences doi: 10.1098/rspb.2009.0941   

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Hierarchical properties characterize elephant fission–fusion social organization whereby stable groups of individuals coalesce into higher order groups or split in a predictable manner. This hierarchical complexity is rare among animals and, as such, an examination of the factors driving its emergence offers unique insight into the evolution of social behaviour. Investigation of the genetic basis for such social affiliation demonstrates that while the majority of core social groups (second-tier affiliates) are significantly related, this is not exclusively the case. As such, direct benefits received through membership of these groups appear to be salient to their formation and maintenance. Further analysis revealed that the majority of groups in the two higher social echelons (third and fourth tiers) are typically not significantly related.

Wittemyer, G., Polansky, L., Douglas-Hamilton, I., Getz, W.M.
Disentangling the effects of forage, social rank and risk on movement autocorrelation of elephants using Fourier and wavelet analysis. (2008)
Proceedings of the National Academy of Sciences 105:19108-19113

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The internal state of an individual—as it relates to thirst, hunger, fear, or reproductive drive—can be inferred by referencing points on its movement path to external environmental and sociological variables. Using time-series approaches to characterize autocorrelative properties of step-length movements collated every 3 h for seven free-ranging African elephants, we examined the influence of social rank, predation risk, and seasonal variation in resource abundance on periodic properties of movement. The frequency domain methods of Fourier and wavelet analyses provide compact summaries of temporal autocorrelation and show both strong diurnal and seasonal based periodicities in the step-length time series. This autocorrelation is weaker during the wet season, indicating random movements are more common when ecological conditions are good.

Wittemyer, G., Rasmussen, H.B., Douglas-Hamilton, I
Breeding phenology in relation to NDVI variability in free-ranging African elephant (2007)
Ecography 30: 42-50, 2007

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The phenology of reproduction is often correlated with resource availability and is hypothesized to be shaped by selective forces in order to maximize lifetime reproductive success. African elephants have the distinctive life history traits of a 22 month gestation and extended offspring investment, necessitating a long-term strategy of energy acquisition and reproductive expenditure to ensure successful offspring recruitment. We investigated the relationship between the reproductive phenology of a wild elephant population and resource availability using remotely sensed Normalized differential vegetation index (NDVI) data as a measure of time-specific primary productivity and hence forage quality.