BackgroundThe increasing human population and global intensification of agriculture have had a major impact on the world’s natural ecosystems and caused devastating effects on populations of mega-herbivores such as the African savanna elephants, through habitat reduction and fragmentation and increased human–animal conflict. Animals with vast home ranges are forced into increasingly smaller geographical areas, often restricted by fencing or encroaching anthropogenic activities, resulting in huge pressures on these areas to meet the animals’ resource needs. This can present a nutritional challenge and cause animals to adapt their movement patterns to meet their dietary needs for specific minerals, potentially causing human–animal conflict. The aim of this review is to consolidate understanding of nutritional drivers for animal movement, especially that of African savanna elephants and focus the direction of future research. Peer reviewed literature available was generally geographically specific and studies conducted on isolated populations of individual species. African savanna elephants have the capacity to extensively alter the landscape and have been more greatly studied than other herbivores, making them a good example species to use for this review. Alongside this, their movement choices, potentially linked with nutritional drivers could be applicable to a range of other species. Relevant case study examples of other herbivores moving based on nutritional needs are discussed.MethodsThree databases were searched in this review: Scopus, Web of Science and Google Scholar, using identified search terms. Inclusion and exclusion criteria were determined and applied as required. Additional grey literature was reviewed as appropriate.ResultsInitial searches yielded 1,870 records prior to application of inclusion and exclusion criteria. A less detailed review of grey literature, and additional peer-reviewed literature which did not meet the inclusion criteria but was deemed relevant by the authors was also conducted to ensure thorough coverage of the subject.DiscussionA review of peer reviewed literature was undertaken to examine nutritional drivers for African elephant movement, exploring documented examples from free-ranging African savanna elephants and, where relevant, other herbivore species. This could help inform prediction or mitigation of human–elephant conflict, potentially when animals move according to nutritional needs, and related drivers for this movement. In addition, appropriate grey literature was included to capture current research.

Background: Culicoides biting midges are biological vectors of arboviruses including bluetongue virus (BTV), Schmallenberg virus (SBV) and African horse sickness virus (AHSV). Zoos are home to a wide range of 'at risk' exotic and native species of animals. These animals have a high value both in monetary terms, conservation significance and breeding potential. To understand the risk these viruses pose to zoo animals, it is necessary to characterise the Culicoides fauna at zoos and determine which potential vector species are feeding on which hosts. Methods: Light-suction traps were used at two UK zoos: the Zoological Society of London (ZSL) London Zoo (LZ) and ZSL Whipsnade Zoo (WZ). Traps were run one night each week from June 2014 to June 2015. Culicoides were morphologically identified to the species level and any blood-fed Culicoides were processed for blood-meal analysis. DNA from blood meals was extracted and amplified using previously published primers. Sequencing was then carried out to determine the host species. Results: A total of 11,648 Culicoides were trapped and identified (n = 5880 from ZSL WZ; n = 5768 from ZSL LZ), constituting 25 different species. The six putative vectors of BTV, SBV and AHSV in northern Europe were found at both zoos and made up the majority of the total catch (n = 10,701). A total of 31 host sequences were obtained from blood-fed Culicoides. Culicoides obsoletus/C. scoticus, Culicoides dewulfi, Culicoides parroti and Culicoides punctatus were found to be biting a wide range of mammals including Bactrian camels, Indian rhinoceros, Asian elephants and humans, with Culicoides obsoletus/C. scoticus also biting Darwin's rhea. The bird-biting species, Culicoides achrayi, was found to be feeding on blackbirds, blue tits, magpies and carrion crows. Conclusions: To our knowledge, this is the first study to directly confirm blood-feeding of Culicoides on exotic zoo animals in the UK and shows that they are able to utilise a wide range of exotic as well as native host species. Due to the susceptibility of some zoo animals to Culicoides-borne arboviruses, this study demonstrates that in the event of an outbreak of one of these viruses in the UK, preventative and mitigating measures would need to be taken.

Historically, behaviour of zoo housed species during hours of limited staff and visitor presence has been poorly studied, largely due to the lack of appropriate technology. Advances in digital monitoring equipment and facility design by European elephant holders has given researchers scope to accurately evaluate behaviour for this species over 24 hrs. Various behavioural indicators of welfare have now been identified for zoo housed elephants; however the relationship between resting behaviour and welfare experience has been an area highlighted to require additional research. Lying rest is a potential positive welfare indicator for this species, with studies suggesting that engagement in lying rest can be used to monitor both psychological and physiological wellbeing. Throughout this work we aim to give insights into the behaviour of individual Asian elephants at ZSL Whipsnade Zoo, specifically between the hours of 16:00 and 10:00. In addition to presenting the activity budgets of our study individuals during these times, we explore individual engagement in resting behaviour. Furthermore, we evaluate the social associations of our study group during rest. We provide evidence that unrelated individuals can form strong associations with conspecifics when resting and show that life history is a factor to consider when evaluating social compatibility between group members. Finally, we demonstrate the positive role that calves and juvenile individuals can play in facilitating meaningful associations between group members during rest. Our study highlights the importance of evaluating behaviour during understudied time periods in order to obtain a holistic view of individual welfare, further emphasising the importance of adopting an evidence-based approach to management for this species in zoos.

In June 2017, the Secretary of State's Standards of Modern Zoo Practice (SSSMZP) were updated with an appendix relating specifically to elephants (Appendix 8.8: Elephants). This update was published to bring elephant management standards in line with recognized advancing best practice. All zoos in the UK holding elephants are inspected against the new appendix, in accordance with the Zoo Licensing Act 1981, by dedicated Department for Environment, Food and Rural Affairs (Defra)‐appointed inspectors. Achieving the standards set out within the new appendix will require financial investment and careful planning from all the zoos holding elephants within the UK. At the time of writing, the annual cost of keeping a breeding group of elephants at ZSL Whipsnade Zoo, UK, was calculated from data collected over the last 10 years and the financial investment required to achieve SSSMZP compliance was estimated. The commercial benefit that elephants bring to ZSL Whipsnade Zoo was also quantified using feedback from visitor surveys. The cost of keeping a breeding herd of elephants at ZSL Whipsnade Zoo was estimated at £593 021–£641 863 per year, excluding indirect staffing costs, ground rent and contributions made by the Zoological Society of London (ZSL) to field‐conservation projects. Costs for achieving SSSMZP compliance will be considerably greater with substantial capital investment required. The commercial benefit was found to be extensive; with predicted significant increased visitor dwell time and secondary spend in the presence of elephants. Using ZSL Whipsnade Zoo as an example, this paper aims to consider holistically the financial costs when planning and implementing an optimum, welfare‐centred, sustainable future for elephants in zoos.

The unique geochemistry surrounding the Palabora Mining Company (PMC) land may act as a micronutrient hotspot, attracting elephants to the area. The PMC produces refined copper and extracts phosphates and other minerals. Understanding the spatial influence of geochemistry on the home range size of African elephants is important for elephant population management and conservation.The home ranges of collared elephants surrounding the PMC were significantly smaller (P=0.001) than conspecifics in surrounding reserves, suggesting that their resource needs were met within these smaller areas. Environmental samples (soil, water and plants) were analysed from the mine area and along six transects radiating from the mine centre. Tail hair and faecal samples from elephants at the PMC, and conspecifics within the surrounding area were analysed. All samples were analysed for minerals essential to health and potentially toxic elements (PTEs; As,

The aim of this study was twofold: (1) identify suitable bio-indicators to assess elemental status in elephants using captive elephant samples, and (2) understand how geochemistry influences mineral intake. Tail hair, toenail, faeces, plasma and urine were collected quarterly from 21 elephants at five UK zoos. All elephant food, soil from enclosure(s), and drinking water were also sampled. Elemental analysis was conducted on all samples, using inductively coupled plasma mass spectrometry, focusing on biologically functional minerals (Ca, Cu, Fe, K, Mg, Mn, Na, P, Se and Zn) and trace metals (As, Cd, Pb, U and V). Linear mixed modelling was used to identify how keeper-fed diet, water and soil were reflected in sample bio-indicators. No sample matrix reflected the status of all assessed elements. Toenail was the best bio-indicator of intake for the most elements reviewed in this study, with keeperfed diet being the strongest predictor. Calcium status was reflected in faeces, (p 0.019, R 2 between elephant within zoo-0.608). In this study urine was of no value in determining mineral status here and plasma was of limited value. Results aimed to define the most suitable bio-indicators to assess captive animal health and encourage onward application to wildlife management. Formulation of an appropriate zoo diet requires husbandry skills and applied nutritional science 1. Although there is limited agreement in the literature, the use of appropriate bio-indicators to assess elemental status was suggested by Combs et al. 2 to support evidence-based zoo diet assessment. Zoos in the United Kingdom have a responsibility to provide appropriate nutrition to all animals within their care 3 to prevent nutritional-related disease, compromised welfare and potential reproductive failure. Limited information exists for estimated mineral requirements of elephants 4 , with cases of specific mineral deficiency documented. Due to elephants' low growth rate and large size, clinical signs of nutrient deficiency may go unnoticed for long periods of time 5 , making nutritional evaluation challenging. Jansman and Pas (2015) 6 defined mineral status as the balance between dietary intake of a nutrient and its requirement in the body. Twenty-eight "essential" mineral elements have known metabolic roles in the mammalian system, for which dietary deficiency will lead to clinical deficiency. These include calcium (Ca), phosphorus (P), magnesium (Mg), selenium (Se) and zinc (Zn) 7,8. Minerals are utilized within the body in various forms or individual compartments, with a central reserve or interchange compartment, usually blood and one or more storage compartments, usually bone or liver. Element and animal species affects the speed of mobilisation of the mineral between compartment(s) 7,8. Mineral status can also be altered by interactions between dietary components; for example an increase in dietary P causes a decrease in serum Ca 9,10 , and variations in individuals' metabolism, circadian patterns and pathological state. Analysing elemental sta...

Over the last two decades, criticisms were raised regarding the welfare experienced by elephants in European and North American zoos. Concerns regarding the welfare of zoo-housed elephants in the UK and Europe were consolidated in the publication of several key reports, and media interest peaked. Throughout this study we aim to outline the behavioural measures of welfare observed in the current group of Asian elephants (Elephas maximus) at Zoological Society of London (ZSL) Whipsnade Zoo, using key welfare indicators for this species and comparing them to previous published work. Following the instigation of a species-specific research programme, empirical behavioural data were available to quantify any developments in care and welfare. The collection of behavioural information revealed that individuals in our study group engage in low levels of stereotypic behaviour, have formed and maintain strong associations with one another and display a high proportion of engagement in lying rest. We outline that by applying simple, low-cost methods of behavioural data collection and analysis, it is possible to collect evidence that allows us to evaluate individual level welfare. This facilitates the adoption of an evidence-based approach to zoo management as well as demonstrating compliance with updated legislation for this species.

The articles in this volume of the International Zoo Yearbook highlight the contributions that professional zoological institutions have made to the care and welfare of elephants over the last 20 years, and describe the threats faced by these iconic animals in their natural habitats. Zoos work to protect wild elephants using a variety of methods, such as carrying out direct research with ex situ populations, providing conservation education and conservation funding, and engaging in capacity building. Furthermore, important work is implemented by conservation organizations in the field. This volume focuses attention on elephant conservation and the important role played by zoological institutions and other dedicated conservation organizations. It is hoped that this work will inspire more zoos to act, help to inform future conservation initiatives and, ultimately, ensure the survival of these iconic species and the protection of their habitats. (Photo: Robin Winkelman, Saint Louis Zoo)