Interplay of competition and facilitation in grazing succession by migrant Serengeti herbivores.

Competition, facilitation, and predation offer alternative explanations for successional patterns of migratory herbivores. However, these interactions are difficult to measure, leaving uncertainty about the mechanisms underlying body-size-dependent grazing-and even whether succession occurs at all. We used data from an 8-year camera-trap survey, GPS-collared herbivores, and fecal DNA metabarcoding to analyze the timing, arrival order, and interactions among migratory grazers in Serengeti National Park. Temporal grazing succession is characterized by a "push-pull" dynamic: Competitive grazing nudges zebra ahead of co-migrating wildebeest, whereas grass consumption by these large-bodied migrants attracts trailing, small-bodied gazelle that benefit from facilitation. "Natural experiments" involving intense wildfires and rainfall respectively disrupted and strengthened these effects. Our results highlight a balance between facilitative and competitive forces in co-regulating large-scale ungulate migrations.

Using sulfur stable isotope ratios (δ34 S) for animal geolocation: Estimating the delay mechanisms between diet ingestion and isotope incorporation in tail hair.

RATIONALE: Metabolism and diet quality play an important role in determining delay mechanisms between an animal ingesting an element and depositing the associated isotope signal in tissue. While many isotope mixing models assume instantaneous reflection of diet in an animal- tissue, this is rarely the case. Here we use data from wildebeest to measure the lag time between ingestion of 34 S and its detection in tail hair. METHODS: We use time-lagged regression analysis of δ34 S data from GPS-collared blue wildebeest from the Serengeti ecosystem in combination with δ34 S isoscape data to estimate the lag time between an animal ingesting and depositing 34 S in tail hair. RESULTS: The best fitting regression model of δ34 S in tail hair and an individual- position on the δ34 S isoscape is generated assuming an average time delay of 78 days between ingestion and detection in tail hair. This suggests that sulfur may undergo multiple metabolic transitions before being deposited in tissue. CONCLUSION: Our findings help to unravel the underlying complexities associated with sulfur metabolism and are broadly consistent with results from other species. These findings will help to inform research aiming to apply the variation of δ34 S in inert biological material for geolocation or understanding dietary changes, especially for fast moving migratory ungulates such as wildebeest.

Local and wide-scale livestock movement networks inform disease control strategies in East Africa.

Livestock mobility exacerbates infectious disease risks across sub-Saharan Africa, but enables critical access to grazing and water resources, and trade. Identifying locations of high livestock traffic offers opportunities for targeted control. We focus on Tanzanian agropastoral and pastoral communities that account respectively for over 75% and 15% of livestock husbandry in eastern Africa. We construct networks of livestock connectivity based on participatory mapping data on herd movements reported by village livestock keepers as well as data from trading points to understand how seasonal availability of resources, land-use and trade influence the movements of livestock. In communities that practise agropastoralism, inter- and intra-village connectivity through communal livestock resources (e.g. pasture and water) was 1.9 times higher in the dry compared to the wet season suggesting greater livestock traffic and increased contact probability. In contrast, livestock from pastoral communities were 1.6 times more connected at communal locations during the wet season when they also tended to move farther (by 3 km compared to the dry season). Trade-linked movements were twice more likely from rural to urban locations. Urban locations were central to all networks, particularly those with potentially high onward movements, for example to abattoirs, livestock holding grounds, or other markets, including beyond national boundaries. We demonstrate how livestock movement information can be used to devise strategic interventions that target critical livestock aggregation points (i.e. locations of high centrality values) and times (i.e. prior to and after the wet season in pastoral and agropastoral areas, respectively). Such targeted interventions are a cost-effective approach to limit infection without restricting livestock mobility critical to sustainable livelihoods.

Deep learning enables satellite-based monitoring of large populations of terrestrial mammals across heterogeneous landscape.

New satellite remote sensing and machine learning techniques offer untapped possibilities to monitor global biodiversity with unprecedented speed and precision. These efficiencies promise to reveal novel ecological insights at spatial scales which are germane to the management of populations and entire ecosystems. Here, we present a robust transferable deep learning pipeline to automatically locate and count large herds of migratory ungulates (wildebeest and zebra) in the Serengeti-Mara ecosystem using fine-resolution (38-50 cm) satellite imagery. The results achieve accurate detection of nearly 500,000 individuals across thousands of square kilometers and multiple habitat types, with an overall F1-score of 84.75% (Precision: 87.85%, Recall: 81.86%). This research demonstrates the capability of satellite remote sensing and machine learning techniques to automatically and accurately count very large populations of terrestrial mammals across a highly heterogeneous landscape. We also discuss the potential for satellite-derived species detections to advance basic understanding of animal behavior and ecology.

Inferring spatially varying animal movement characteristics using a hierarchical continuous-time velocity model.

Understanding the spatial dynamics of animal movement is an essential component of maintaining ecological connectivity, conserving key habitats, and mitigating the impacts of anthropogenic disturbance. Altered movement and migratory patterns are often an early warning sign of the effects of environmental disturbance, and a precursor to population declines. Here, we present a hierarchical Bayesian framework based on Gaussian processes for analysing the spatial characteristics of animal movement. At the heart of our approach is a novel covariance kernel that links the spatially varying parameters of a continuous-time velocity model with GPS locations from multiple individuals. We demonstrate the effectiveness of our framework by first applying it to a synthetic data set and then by analysing telemetry data from the Serengeti wildebeest migration. Through application of our approach, we are able to identify the key pathways of the wildebeest migration as well as revealing the impacts of environmental features on movement behaviour.

Livestock movement informs the risk of disease spread in traditional production systems in East Africa.

In Africa, livestock are important to local and national economies, but their productivity is constrained by infectious diseases. Comprehensive information on livestock movements and contacts is required to devise appropriate disease control strategies; yet, understanding contact risk in systems where herds mix extensively, and where different pathogens can be transmitted at different spatial and temporal scales, remains a major challenge. We deployed Global Positioning System collars on cattle in 52 herds in a traditional agropastoral system in western Serengeti, Tanzania, to understand fine-scale movements and between-herd contacts, and to identify locations of greatest interaction between herds. We examined contact across spatiotemporal scales relevant to different disease transmission scenarios. Daily cattle movements increased with herd size and rainfall. Generally, contact between herds was greatest away from households, during periods with low rainfall and in locations close to dipping points. We demonstrate how movements and contacts affect the risk of disease spread. For example, transmission risk is relatively sensitive to the survival time of different pathogens in the environment, and less sensitive to transmission distance, at least over the range of the spatiotemporal definitions of contacts that we explored. We identify times and locations of greatest disease transmission potential and that could be targeted through tailored control strategies.

Drivers of site fidelity in ungulates.

While the tendency to return to previously visited locations-termed 'site fidelity'-is common in animals, the cause of this behaviour is not well understood. One hypothesis is that site fidelity is shaped by an animal's environment, such that animals living in landscapes with predictable resources have stronger site fidelity. Site fidelity may also be conditional on the success of animals' recent visits to that location, and it may become stronger with age as the animal accumulates experience in their landscape. Finally, differences between species, such as the way memory shapes site attractiveness, may interact with environmental drivers to modulate the strength of site fidelity. We compared inter-year site fidelity in 669 individuals across eight ungulate species fitted with GPS collars and occupying a range of environmental conditions in North America and Africa. We used a distance-based index of site fidelity and tested hypothesized drivers of site fidelity using linear mixed effects models, while accounting for variation in annual range size. Mule deer Odocoileus hemionus and moose Alces alces exhibited relatively strong site fidelity, while wildebeest Connochaetes taurinus and barren-ground caribou Rangifer tarandus granti had relatively weak fidelity. Site fidelity was strongest in predictable landscapes where vegetative greening occurred at regular intervals over time (i.e. high temporal contingency). Species differed in their response to spatial heterogeneity in greenness (i.e. spatial constancy). Site fidelity varied seasonally in some species, but remained constant over time in others. Elk employed a 'win-stay, lose-switch' strategy, in which successful resource tracking in the springtime resulted in strong site fidelity the following spring. Site fidelity did not vary with age in any species tested. Our results provide support for the environmental hypothesis, particularly that regularity in vegetative phenology shapes the strength of site fidelity at the inter-annual scale. Large unexplained differences in site fidelity suggest that other factors, possibly species-specific differences in attraction to known sites, contribute to variation in the expression of this behaviour. Understanding drivers of variation in site fidelity across groups of organisms living in different environments provides important behavioural context for predicting how animals will respond to environmental change.

Author Correction: Longest terrestrial migrations and movements around the world.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Longest terrestrial migrations and movements around the world.

Long-distance terrestrial migrations are imperiled globally. We determined both round-trip migration distances (straight-line measurements between migratory end points) and total annual movement (sum of the distances between successive relocations over a year) for a suite of large mammals that had potential for long-distance movements to test which species displayed the longest of both. We found that caribou likely do exhibit the longest terrestrial migrations on the planet, but, over the course of a year, gray wolves move the most. Our results were consistent with the trophic-level based hypothesis that predators would move more than their prey. Herbivores in low productivity environments moved more than herbivores in more productive habitats. We also found that larger members of the same guild moved less than smaller members, supporting the 'gastro-centric' hypothesis. A better understanding of migration and movements of large mammals should aid in their conservation by helping delineate conservation area boundaries and determine priority corridors for protection to preserve connectivity. The magnitude of the migrations and movements we documented should also provide guidance on the scale of conservation efforts required and assist conservation planning across agency and even national boundaries.

Transmission ecology of canine parvovirus in a multi-host, multi-pathogen system.

Understanding multi-host pathogen maintenance and transmission dynamics is critical for disease control. However, transmission dynamics remain enigmatic largely because they are difficult to observe directly, particularly in wildlife. Here, we investigate the transmission dynamics of canine parvovirus (CPV) using state-space modelling of 20 years of CPV serology data from domestic dogs and African lions in the Serengeti ecosystem. We show that, although vaccination reduces the probability of infection in dogs, and despite indirect enhancement of population seropositivity as a result of vaccine shedding, the vaccination coverage achieved has been insufficient to prevent CPV from becoming widespread. CPV is maintained by the dog population and has become endemic with approximately 3.5-year cycles and prevalence reaching approximately 80%. While the estimated prevalence in lions is lower, peaks of infection consistently follow those in dogs. Dogs exposed to CPV are also more likely to become infected with a second multi-host pathogen, canine distemper virus. However, vaccination can weaken this coupling, raising questions about the value of monovalent versus polyvalent vaccines against these two pathogens. Our findings highlight the need to consider both pathogen- and host-level community interactions when seeking to understand the dynamics of multi-host pathogens and their implications for conservation, disease surveillance and control programmes.

Cross-boundary human impacts compromise the Serengeti-Mara ecosystem.

Protected areas provide major benefits for humans in the form of ecosystem services, but landscape degradation by human activity at their edges may compromise their ecological functioning. Using multiple lines of evidence from 40 years of research in the Serengeti-Mara ecosystem, we find that such edge degradation has effectively "squeezed" wildlife into the core protected area and has altered the ecosystem's dynamics even within this 40,000-square-kilometer ecosystem. This spatial cascade reduced resilience in the core and was mediated by the movement of grazers, which reduced grass fuel and fires, weakened the capacity of soils to sequester nutrients and carbon, and decreased the responsiveness of primary production to rainfall. Similar effects in other protected ecosystems worldwide may require rethinking of natural resource management outside protected areas.

From single steps to mass migration: the problem of scale in the movement ecology of the Serengeti wildebeest.

A central question in ecology is how to link processes that occur over different scales. The daily interactions of individual organisms ultimately determine community dynamics, population fluctuations and the functioning of entire ecosystems. Observations of these multiscale ecological processes are constrained by various technological, biological or logistical issues, and there are often vast discrepancies between the scale at which observation is possible and the scale of the question of interest. Animal movement is characterized by processes that act over multiple spatial and temporal scales. Second-by-second decisions accumulate to produce annual movement patterns. Individuals influence, and are influenced by, collective movement decisions, which then govern the spatial distribution of populations and the connectivity of meta-populations. While the field of movement ecology is experiencing unprecedented growth in the availability of movement data, there remain challenges in integrating observations with questions of ecological interest. In this article, we present the major challenges of addressing these issues within the context of the Serengeti wildebeest migration, a keystone ecological phenomena that crosses multiple scales of space, time and biological complexity.This article is part of the theme issue 'Collective movement ecology'.

Assessing Rotation-Invariant Feature Classification for Automated Wildebeest Population Counts.

Accurate and on-demand animal population counts are the holy grail for wildlife conservation organizations throughout the world because they enable fast and responsive adaptive management policies. While the collection of image data from camera traps, satellites, and manned or unmanned aircraft has advanced significantly, the detection and identification of animals within images remains a major bottleneck since counting is primarily conducted by dedicated enumerators or citizen scientists. Recent developments in the field of computer vision suggest a potential resolution to this issue through the use of rotation-invariant object descriptors combined with machine learning algorithms. Here we implement an algorithm to detect and count wildebeest from aerial images collected in the Serengeti National Park in 2009 as part of the biennial wildebeest count. We find that the per image error rates are greater than, but comparable to, two separate human counts. For the total count, the algorithm is more accurate than both manual counts, suggesting that human counters have a tendency to systematically over or under count images. While the accuracy of the algorithm is not yet at an acceptable level for fully automatic counts, our results show this method is a promising avenue for further research and we highlight specific areas where future research should focus in order to develop fast and accurate enumeration of aerial count data. If combined with a bespoke image collection protocol, this approach may yield a fully automated wildebeest count in the near future.

Balancing Conservation with National Development: A Socio-Economic Case Study of the Alternatives to the Serengeti Road.

Developing countries often have rich natural resources but poor infrastructure to capitalize on them, which leads to significant challenges in terms of balancing poverty alleviation with conservation. The underlying premise in development strategies is to increase the socio-economic welfare of the people while simultaneously ensuring environmental sustainability, however these objectives are often in direct conflict. National progress is dependent on developing infrastructure such as effective transportation networks, however roads can be ecologically catastrophic in terms of disrupting habitat connectivity and facilitating illegal activity. How can national development and conservation be balanced? The proposed Serengeti road epitomizes the conflict between poverty alleviation on one hand, and the conservation of a critical ecosystem on the other. We use the Serengeti as an exemplar case-study in which the relative economic and social benefits of a road can be assessed against the ecological impacts. Specifically, we compare three possible transportation routes and ask which route maximizes the socio-economic returns for the people while minimizing the ecological costs. The findings suggest that one route in particular that circumnavigates the Serengeti links the greatest number of small and medium sized entrepreneurial businesses to the largest labour force in the region. Furthermore, this route connects the most children to schools, provisions the greatest access to hospitals, and opens the most fertile crop and livestock production areas, and does not compromise the ecology and tourism revenue of the Serengeti. This route would improve Tanzania's food security and self-reliance and would facilitate future infrastructure development which would not be possible if the road were to pass through the Serengeti. This case study provides a compelling example of how a detailed spatial analysis can balance the national objectives of poverty alleviation while maintaining ecological integrity.

Dynamics of a morbillivirus at the domestic-wildlife interface: Canine distemper virus in domestic dogs and lions.

Morbilliviruses cause many diseases of medical and veterinary importance, and although some (e.g., measles and rinderpest) have been controlled successfully, others, such as canine distemper virus (CDV), are a growing concern. A propensity for host-switching has resulted in CDV emergence in new species, including endangered wildlife, posing challenges for controlling disease in multispecies communities. CDV is typically associated with domestic dogs, but little is known about its maintenance and transmission in species-rich areas or about the potential role of domestic dog vaccination as a means of reducing disease threats to wildlife. We address these questions by analyzing a long-term serological dataset of CDV in lions and domestic dogs from Tanzania's Serengeti ecosystem. Using a Bayesian state-space model, we show that dynamics of CDV have changed considerably over the past three decades. Initially, peaks of CDV infection in dogs preceded those in lions, suggesting that spill-over from dogs was the main driver of infection in wildlife. However, despite dog-to-lion transmission dominating cross-species transmission models, infection peaks in lions became more frequent and asynchronous from those in dogs, suggesting that other wildlife species may play a role in a potentially complex maintenance community. Widespread mass vaccination of domestic dogs reduced the probability of infection in dogs and the size of outbreaks but did not prevent transmission to or peaks of infection in lions. This study demonstrates the complexity of CDV dynamics in natural ecosystems and the value of long-term, large-scale datasets for investigating transmission patterns and evaluating disease control strategies.

African vultures don't follow migratory herds: scavenger habitat use is not mediated by prey abundance.

The ongoing global decline in vulture populations raises major conservation concerns, but little is known about the factors that mediate scavenger habitat use, in particular the importance of abundance of live prey versus prey mortality. We test this using data from the Serengeti-Mara ecosystem in East Africa. The two hypotheses that prey abundance or prey mortality are the main drivers of vulture habitat use provide alternative predictions. If vultures select areas based only on prey abundance, we expect tracked vultures to remain close to herds of migratory wildebeest regardless of season. However, if vultures select areas where mortality rates are greatest then we expect vultures to select the driest regions, where animals are more likely to die of starvation, and to be attracted to migratory wildebeest only during the dry season when wildebeest mortality is greatest. We used data from GSM-GPS transmitters to assess the relationship between three vulture species and migratory wildebeest in the Mara-Serengeti ecosystem. Results indicate that vultures preferentially cluster around migratory herds only during the dry season, when herds experience their highest mortality. Additionally during the wet season, Ruppell's and Lappet-faced vultures select relatively dry areas, based on Normalized Difference Vegetation Index, whereas White-backed vultures preferred wetter areas during the wet season. Differences in habitat use among species may mediate coexistence in this scavenger guild. In general, our results suggest that prey abundance is not the primary driver of avian scavenger habitat use. The apparent reliance of vultures on non-migratory ungulates during the wet season has important conservation implications for vultures in light of on-going declines in non-migratory ungulate species and use of poisons in unprotected areas.

Evaluating the potential for the environmentally sustainable control of foot and mouth disease in Sub-Saharan Africa.

Strategies to control transboundary diseases have in the past generated unintended negative consequences for both the environment and local human populations. Integrating perspectives from across disciplines, including livestock, veterinary and conservation sectors, is necessary for identifying disease control strategies that optimise environmental goods and services at the wildlife-livestock interface. Prompted by the recent development of a global strategy for the control and elimination of foot-and-mouth disease (FMD), this paper seeks insight into the consequences of, and rational options for potential FMD control measures in relation to environmental, conservation and human poverty considerations in Africa. We suggest a more environmentally nuanced process of FMD control that safe-guards the integrity of wild populations and the ecosystem dynamics on which human livelihoods depend while simultaneously improving socio-economic conditions of rural people. In particular, we outline five major issues that need to be considered: 1) improved understanding of the different FMD viral strains and how they circulate between domestic and wildlife populations; 2) an appreciation for the economic value of wildlife for many African countries whose presence might preclude the country from ever achieving an FMD-free status; 3) exploring ways in which livestock production can be improved without compromising wildlife such as implementing commodity-based trading schemes; 4) introducing a participatory approach involving local farmers and the national veterinary services in the control of FMD; and 5) finally the possibility that trans frontier conservation might offer new hope of integrating decision-making at the wildlife-livestock interface.