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BackgroundMovement is central to understanding the ecology of animals. The most robustly definable segments of an individual's lifetime track are its diel activity routines (DARs). This robustness is due to fixed start and end points set by a 24-h clock that depends on the individual's quotidian schedule. An analysis of day-to-day variation in the DARs of individuals, their comparisons among individuals, and the questions that can be asked, particularly in the context of lunar and annual cycles, depends on the relocation frequency and spatial accuracy of movement data. Here we present methods for categorizing the geometry of DARs for high frequency (seconds to minutes) movement data.MethodsOur method involves an initial categorization of DARs using data pooled across all individuals. We approached this categorization using a Ward clustering algorithm that employs four scalar "whole-path metrics" of trajectory geometry: 1. net displacement (distance between start and end points), 2. maximum displacement from start point, 3. maximum diameter, and 4. maximum width. We illustrate the general approach using reverse-GPS data obtained from 44 barn owls, Tyto alba, in north-eastern Israel. We conducted a principle components analysis (PCA) to obtain a factor, PC1, that essentially captures the scale of movement. We then used a generalized linear mixed model with PC1 as the dependent variable to assess the effects of age and sex on movement.ResultsWe clustered 6230 individual DARs into 7 categories representing different shapes and scale of the owls nightly routines. Five categories based on size and elongation were classified as closed (i.e. returning to the same roost), one as partially open (returning to a nearby roost) and one as fully open (leaving for another region). Our PCA revealed that the DAR scale factor, PC1, accounted for 86.5% of the existing variation. It also showed that PC2 captures the openness of the DAR and accounted for another 8.4% of the variation. We also constructed spatio-temporal distributions of DAR types for individuals and groups of individuals aggregated by age, sex, and seasonal quadrimester, as well as identify some idiosyncratic behavior of individuals within family groups in relation to location. Finally, we showed in two ways that DARs were significantly larger in young than adults and in males than females.ConclusionOur study offers a new method for using high-frequency movement data to classify animal diel movement routines. Insights into the types and distributions of the geometric shape and size of DARs in populations may well prove to be more invaluable for predicting the space-use response of individuals and populations to climate and land-use changes than other currently used movement track methods of analysis.

Resource partitioning promotes coexistence among guild members, and carnivores reduce interference competition through behavioral mechanisms that promote spatio-temporal separation. We analyzed sympatric lion and spotted hyena movements and activity patterns to ascertain the mechanisms facilitating their coexistence within semi-arid and wetland ecosystems. We identified recurrent high-use (revisitation) and extended stay (duration) areas within home ranges, as well as correlated movement-derived measures of inter- and intraspecific interactions with environmental variables. Spatial overlaps among lions and hyenas expanded during the wet season, and occurred at edges of home ranges, around water-points, along pathways between patches of high-use areas. Lions shared more of their home ranges with spotted hyenas in arid ecosystems, but shared more of their ranges with conspecifics in mesic environments. Despite shared space use, we found evidence for subtle temporal differences in the nocturnal movement and activity patterns between the two predators, suggesting a fine localized-scale avoidance strategy. Revisitation frequency and duration within home ranges were influenced by interspecific interactions, after land cover categories and diel cycles. Intraspecific interactions were also important for lions and, important for hyenas were moon illumination and ungulates attracted to former anthrax carcass sites in Etosha, with distance to water in Chobe/Linyanti. Recursion and duration according to locales of competitor probabilities were similar among female lions and both sexes of hyenas, but different for male lions. Our results suggest that lions and spotted hyenas mediate the potential for interference competition through subtle differences in temporal activity, fine-scale habitat use differentiation, and localized reactive-avoidance behaviors. These findings enhance our understanding of the potential effects of interspecific interactions among large carnivore space-use patterns within an apex predator system and show adaptability across heterogeneous and homogeneous environments. Future conservation plans should emphasize the importance of inter- and intraspecific competition within large carnivore communities, particularly moderating such effects within increasingly fragmented landscapes.

We investigated the initial outbreak rates and subsequent social distancing behaviour over the initial phase of the COVID-19 pandemic across 29 Combined Statistical Areas (CSAs) of the United States. We used the Numerus Model Builder Data and Simulation Analysis (NMB-DASA) web application to fit the exponential phase of a SCLAIV+D (Susceptible, Contact, Latent, Asymptomatic infectious, symptomatic Infectious, Vaccinated, Dead) disease classes model to outbreaks, thereby allowing us to obtain an estimate of the basic reproductive number R0 for each CSA. Values of R0 ranged from 1.9 to 9.4, with a mean and standard deviation of 4.5±1.8. Fixing the parameters from the exponential fit, we again used NMB-DASA to estimate a set of social distancing behaviour parameters to compute an epidemic flattening index cflatten. Finally, we applied hierarchical clustering methods using this index to divide CSA outbreaks into two clusters: those presenting a social distancing response that was either weaker or stronger. We found cflatten to be more influential in the clustering process than R0. Thus, our results suggest that the behavioural response after a short initial exponential growth phase is likely to be more determinative of the rise of an epidemic than R0 itself.

BackgroundAn understanding of epidemiological dynamics, once confined to mathematical epidemiologists and applied mathematicians, can be disseminated to a non-mathematical community of health care professionals and applied biologists through simple-to-use simulation applications. We used Numerus Model Builder RAMP Ⓡ (Runtime Alterable Model Platform) technology, to construct deterministic and stochastic versions of compartmental SIR (Susceptible, Infectious, Recovered with immunity) models as simple-to-use, freely available, epidemic simulation application programs.ResultsWe take the reader through simulations used to demonstrate the following concepts: 1) disease prevalence curves of unmitigated outbreaks have a single peak and result in epidemics that 'burn' through the population to become extinguished when the proportion of the susceptible population drops below a critical level; 2) if immunity in recovered individuals wanes sufficiently fast then the disease persists indefinitely as an endemic state, with possible dampening oscillations following the initial outbreak phase; 3) the steepness and initial peak of the prevalence curve are influenced by the basic reproductive value R0, which must exceed 1 for an epidemic to occur; 4) the probability that a single infectious individual in a closed population (i.e. no migration) gives rise to an epidemic increases with the value of R0>1; 5) behavior that adaptively decreases the contact rate among individuals with increasing prevalence has major effects on the prevalence curve including dramatic flattening of the prevalence curve along with the generation of multiple prevalence peaks; 6) the impacts of treatment are complicated to model because they effect multiple processes including transmission, recovery and mortality; 7) the impacts of vaccination policies, constrained by a fixed number of vaccination regimens and by the rate and timing of delivery, are crucially important to maximizing the ability of vaccination programs to reduce mortality.ConclusionOur presentation makes transparent the key assumptions underlying SIR epidemic models. Our RAMP simulators are meant to augment rather than replace classroom material when teaching epidemiological dynamics. They are sufficiently versatile to be used by students to address a range of research questions for term papers and even dissertations.

BackgroundNo versatile web app exists that allows epidemiologists and managers around the world to comprehensively analyze the impacts of COVID-19 mitigation. The http://covid-webapp.numerusinc.com/ web app presented here fills this gap.MethodsOur web app uses a model that explicitly identifies susceptible, contact, latent, asymptomatic, symptomatic and recovered classes of individuals, and a parallel set of response classes, subject to lower pathogen-contact rates. The user inputs a CSV file of incidence and, if of interest, mortality rate data. A default set of parameters is available that can be overwritten through input or online entry, and a user-selected subset of these can be fitted to the model using maximum-likelihood estimation (MLE). Model fitting and forecasting intervals are specifiable and changes to parameters allow counterfactual and forecasting scenarios. Confidence or credible intervals can be generated using stochastic simulations, based on MLE values, or on an inputted CSV file containing Markov chain Monte Carlo (MCMC) estimates of one or more parameters.ResultsWe illustrate the use of our web app in extracting social distancing, social relaxation, surveillance or virulence switching functions (i.e., time varying drivers) from the incidence and mortality rates of COVID-19 epidemics in Israel, South Africa, and England. The Israeli outbreak exhibits four distinct phases: initial outbreak, social distancing, social relaxation, and a second wave mitigation phase. An MCMC projection of this latter phase suggests the Israeli epidemic will continue to produce into late November an average of around 1500 new case per day, unless the population practices social-relaxation measures at least 5-fold below the level in August, which itself is 4-fold below the level at the start of July. Our analysis of the relatively late South African outbreak that became the world's fifth largest COVID-19 epidemic in July revealed that the decline through late July and early August was characterised by a social distancing driver operating at more than twice the per-capita applicable-disease-class (pc-adc) rate of the social relaxation driver. Our analysis of the relatively early English outbreak, identified a more than 2-fold improvement in surveillance over the course of the epidemic. It also identified a pc-adc social distancing rate in early August that, though nearly four times the pc-adc social relaxation rate, appeared to barely contain a second wave that would break out if social distancing was further relaxed.ConclusionOur web app provides policy makers and health officers who have no epidemiological modelling or computer coding expertise with an invaluable tool for assessing the impacts of different outbreak mitigation policies and measures. This includes an ability to generate an epidemic-suppression or curve-flattening index that measures the intensity with which behavioural responses suppress or flatten the epidemic curve in the region under consideration.

We present methods for building a Java Runtime-Alterable-Model Platform (RAMP) of complex dynamical systems. We illustrate our methods by building a multivariant SEIR (epidemic) RAMP. Underlying our RAMP is an individual-based model that includes adaptive contact rates, pathogen genetic drift, waning and cross-immunity. Besides allowing parameter values, process descriptions and scriptable runtime drivers to be easily modified during simulations, our RAMP can used within R-Studio and other computational platforms. Process descriptions that can be runtime altered within our SEIR RAMP include pathogen variant-dependent host shedding, environmental persistence, host transmission and within-host pathogen mutation and replication. They also include adaptive social distancing and adaptive application of vaccination rates and variant-valency of vaccines. We present simulation results using parameter values and process descriptions relevant to the current COVID-19 pandemic. Our results suggest that if waning immunity outpaces vaccination rates, then vaccination rollouts may fail to contain the most transmissible variants, particularly if vaccine valencies are not adapted to deal with escape mutations. Our SEIR RAMP is designed for easy use by others. More generally, our RAMP concept facilitates construction of highly flexible complex systems models of all types, which can then be easily shared as stand-alone application programs.

When a transmission hotspot for an environmentally persistent pathogen establishes in otherwise high-quality habitat, the disease may exert a strong impact on a host population. However, fluctuating environmental conditions lead to heterogeneity in habitat quality and animal habitat preference, which may interrupt the overlap between selected and risky habitats. We evaluated spatio-temporal patterns in anthrax mortalities in a plains zebra (Equus quagga) population in Etosha National Park, Namibia, incorporating remote-sensing and host telemetry data. A higher proportion of anthrax mortalities of herbivores was detected in open habitats than in other habitat types. Resource selection functions showed that the zebra population shifted habitat selection in response to changes in rainfall and vegetation productivity. Average to high rainfall years supported larger anthrax outbreaks, with animals congregating in preferred open habitats, while a severe drought forced animals into otherwise less preferred habitats, leading to few anthrax mortalities. Thus, the timing of anthrax outbreaks was congruent with preference for open plains habitats and a corresponding increase in pathogen exposure. Given shifts in habitat preference, the overlap in high-quality habitat and high-risk habitat is intermittent, reducing the adverse consequences for the population.

Severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2) the causal agent for COVID-19, is a communicable disease spread through close contact. It is known to disproportionately impact certain communities due to both biological susceptibility and inequitable exposure. In this study, we investigate the most important health, social, and environmental factors impacting the early phases (before July, 2020) of per capita COVID-19 transmission and per capita all-cause mortality in US counties. We aggregate county-level physical and mental health, environmental pollution, access to health care, demographic characteristics, vulnerable population scores, and other epidemiological data to create a large feature set to analyze per capita COVID-19 outcomes. Because of the high-dimensionality, multicollinearity, and unknown interactions of the data, we use ensemble machine learning and marginal prediction methods to identify the most salient factors associated with several COVID-19 outbreak measure. Our variable importance results show that measures of ethnicity, public transportation and preventable diseases are the strongest predictors for both per capita COVID-19 incidence and mortality. Specifically, the CDC measures for minority populations, CDC measures for limited English, and proportion of Black- and/or African-American individuals in a county were the most important features for per capita COVID-19 cases within a month after the pandemic started in a county and also at the latest date examined. For per capita all-cause mortality at day 100 and total to date, we find that public transportation use and proportion of Black- and/or African-American individuals in a county are the strongest predictors. The methods predict that, keeping all other factors fixed, a 10% increase in public transportation use, all other factors remaining fixed at the observed values, is associated with increases mortality at day 100 of 2012 individuals (95% CI [1972, 2356]) and likewise a 10% increase in the proportion of Black- and/or African-American individuals in a county is associated with increases total deaths at end of study of 2067 (95% CI [1189, 2654]). Using data until the end of study, the same metric suggests ethnicity has double the association as the next most important factors, which are location, disease prevalence, and transit factors. Our findings shed light on societal patterns that have been reported and experienced in the U.S. by using robust methods to understand the features most responsible for transmission and sectors of society most vulnerable to infection and mortality. In particular, our results provide evidence of the disproportionate impact of the COVID-19 pandemic on minority populations. Our results suggest that mitigation measures, including how vaccines are distributed, could have the greatest impact if they are given with priority to the highest risk communities.

Escape mutations (EM) to SARS-Cov-2 have been detected and are spreading. Vaccines may need adjustment to respond to these or future mutations. We designed a population level model integrating both waning immunity and EM. We also designed a set of criteria for elaborating and fitting this model to cross-neutralization and other data in a manner that minimizes vaccine decision errors. We formulated four model variations. These define criteria for which prior infections provide immunity that can be escaped. They also specify different sequences where one EM follows another. At all reasonable parameter values, these model variations led to patterns where: 1) EM were rare in the first epidemic, 2) rebound epidemics after the first epidemic were accelerated more by increasing drifting than by increasing waning (with some exceptions), 3) the long term endemic level of infection was determined mostly by waning rates with small effects of the drifting parameter, 4) EM caused loss of vaccine effectiveness and under some conditions, vaccines induced EM that caused higher levels of infection with vaccines than without them. The differences and similarities across the four models suggest paths for developing models specifying the epitopes where EM act. This model is a base on which to construct epitope specific evolutionary models using new high-throughput assay data from population samples to guide vaccine decisions. Highlights This model is the first to integrate both antigenic drifting from escape mutations and immunity waning in continuous time. Tiny amounts of only waning or only escape mutation drifting have small or no effects. Together, they have large effects. There are no or few escape mutations during the first epidemic peak and no effect of drifting parameters on the size of that wave. After the first epidemic peak, escape mutations accumulate rapidly. They increase with increases in waning rates and with increases in the drifting rate. Escape mutations then amplify other escape mutations since these raise the frequency of reinfections. Escape mutations can completely negate the effects of vaccines and even lead to more infections with vaccination than without, especially at very low waning rates. The model generates population level cross-neutralization patterns that enable the model to be fitted to population level serological data. The model can be modified to use laboratory data that determine the epitope specific effects of mutations on ACE2 attachment strength or escape from antibody effects. The model, although currently unable to predict the effects of escape mutations in the real world, opens up a path that can guide model incorporation of molecularly studied escape mutations and improve predictive value. We describe that path. Model analysis indicates that vaccine trials and serological surveys are needed now to detect the effects of epitope specific escape mutations that could cause the loss of vaccine efficacy.

Environmentally mediated indirect pathogen transmission is linked to host movement and foraging in areas where pathogens are maintained in the environment. In the case of anthrax, spores of the causative bacterium Bacillus anthracis are released into the environment following host death and create locally infectious zones (LIZs) around carcass sites; by grazing at LIZs, herbivores are potentially exposed to spores. Here, we used camera traps to assess how ungulate species use carcass sites in southwestern Montana and evaluated how these behaviours may promote indirect anthrax transmission, thus providing, to our knowledge, the first detailed documentation and study of the fine-scale mechanisms underlying foraging-based disease transmission in this ecosystem. We found that carcasses at LIZs significantly increased aboveground biomass of vegetation and concentrations of sodium and phosphorus, potentially making these sites more appealing to grazers. Host behavioural responses to LIZs varied depending on species, sex, season and carcass age; but, overall, our results demonstrated that carcasses or carcass sites serve as an attractant to herbivores in this system. Attraction to LIZs probably represents an increased risk of exposure to B. anthracis and, consequently, increased anthrax transmission rates. Accordingly, continued anthrax surveillance and control strategies are critical in this system.

The behavioural ecology of host species is likely to affect their microbial communities, because host sex, diet, physiology, and movement behaviour could all potentially influence their microbiota. We studied a wild population of barn owls (Tyto alba) and collected data on their microbiota, movement, diet, size, coloration, and reproduction. The composition of bacterial species differed by the sex of the host and female owls had more diverse bacterial communities than their male counterparts. The abundance of two families of bacteria, Actinomycetaceae and Lactobacillaceae, also varied between the sexes, potentially as a result of sex differences in hormones and immunological function, as has previously been found with Lactobacillaceae in the microbiota of mice. Male and female owls did not differ in the prey they brought to the nest, which suggests that dietary differences are unlikely to underlie the differences in their microbiota. The movement behaviour of the owls was associated with the host microbiota in both males and females because owls that moved further from their nest each day had more diverse bacterial communities than owls that stayed closer to their nests. This novel result suggests that the movement ecology of hosts can impact their microbiota, potentially on the basis of their differential encounters with new bacterial species as the hosts move and forage across the landscape. Overall, we found that many aspects of the microbial community are correlated with the behavioural ecology of the host and that data on the microbiota can aid in generating new hypotheses about host behaviour.

Concern for megafauna is increasing among scientists and non-scientists. Many studies have emphasized that megafauna play prominent ecological roles and provide important ecosystem services to humanity. But, what precisely are 'megafauna'? Here, we critically assess the concept of megafauna and propose a goal-oriented framework for megafaunal research. First, we review definitions of megafauna and analyse associated terminology in the scientific literature. Second, we conduct a survey among ecologists and palaeontologists to assess the species traits used to identify and define megafauna. Our review indicates that definitions are highly dependent on the study ecosystem and research question, and primarily rely on ad hoc size-related criteria. Our survey suggests that body size is crucial, but not necessarily sufficient, for addressing the different applications of the term megafauna. Thus, after discussing the pros and cons of existing definitions, we propose an additional approach by defining two function-oriented megafaunal concepts: 'keystone megafauna' and 'functional megafauna', with its variant 'apex megafauna'. Assessing megafauna from a functional perspective could challenge the perception that there may not be a unifying definition of megafauna that can be applied to all eco-evolutionary narratives. In addition, using functional definitions of megafauna could be especially conducive to cross-disciplinary understanding and cooperation, improvement of conservation policy and practice, and strengthening of public perception. As megafaunal research advances, we encourage scientists to unambiguously define how they use the term 'megafauna' and to present the logic underpinning their definition.

The risk of a zoonotic pandemic disease threatens hundreds of millions of people. Emerging infectious diseases also threaten livestock and wildlife populations around the world and can lead to devastating economic damages. China and the USA-due to their unparalleled resources, widespread engagement in activities driving emerging infectious diseases and national as well as geopolitical imperatives to contribute to global health security-play an essential role in our understanding of pandemic threats. Critical to efforts to mitigate risk is building upon existing investments in global capacity to develop training and research focused on the ecological factors driving infectious disease spillover from animals to humans. International cooperation, particularly between China and the USA, is essential to fully engage the resources and scientific strengths necessary to add this ecological emphasis to the pandemic preparedness strategy. Here, we review the world's current state of emerging infectious disease preparedness, the ecological and evolutionary knowledge needed to anticipate disease emergence, the roles that China and the USA currently play as sources and solutions to mitigating risk, and the next steps needed to better protect the global community from zoonotic disease.

We formulate a refined SEIR epidemic model that explicitly includes a contact class C that either thwarts pathogen invasion and returns to the susceptible class S or progresses successively through latent, asymptomatic, and symptomatic classes L, A, and I. Individuals in both A and I may go directly to an immune class V, and in I to a dead class D. We extend this SCLAIV formulation by including a set of drivers that can be used to develop policy to manage current Covid-19 and similar type disease outbreaks. These drivers include surveillance, social distancing (rate and efficacy), social relaxation, quarantining (linked to contact tracing), patient treatment/isolation and vaccination processes, each of which can be represented by a non-negative constant or an s-shaped switching flow. The latter are defined in terms of onset and switching times, initial and final values, and abruptness of switching. We built a Covid-19NMB-DASA web app to generate both deterministic and stochastic solutions to our SCLAIV and drivers model and use incidence and mortality data to provide both maximum-likelihood estimation (MLE) and Bayesian MCMC fitting of parameters. In the context of South African and English Covid-19 incidence data we demonstrate how to both identify and evaluate the role of drivers in ongoing outbreaks. In particular, we show that early social distancing in South Africa likely averted around 80,000 observed cases (actual number is double if only half the cases are observed) during the months of June and July. We also demonstrated that incidence rates in South Africa will increase to between a conservative estimate of 15 and 30 thousand observed cases per day (at a 50% surveillance level) by the end of August if stronger social distancing measures are not effected during July and August, 2020. On different a note, we show that comparably good local MLE fits of the English data using surveillance, social distancing and social relaxation drivers can represent very different kinds of outbreaks—one with close to 90% and another with under 8% immune individuals. This latter result provides a cautionary tale of why fitting SEIR-like models to incidence or prevalence data can be extremely problematic when not anchored by other critical measures, such as levels of immunity in the population. Our presentation illustrates how our SCLAIV formulation can be used to carry out forensic and scenario analyses of disease outbreaks such as Covid-19 in well defined regions.

BackgroundUnderstanding rhino movement behavior, especially their recursive movements, holds significant promise for enhancing rhino conservation efforts, and protecting their habitats and the biodiversity they support. Here we investigate the daily, biweekly, and seasonal recursion behavior of rhinos, to aid conservation applications and increase our foundational knowledge about these important ecosystem engineers.MethodsUsing relocation data from 59 rhinos across northern Namibia and 8 years of sampling efforts, we investigated patterns in 24-h displacement at dawn, dusk, midday, and midnight to examine movement behaviors at an intermediate scale and across daily behavioral modes of foraging and resting. To understand recursion patterns across animals' short and long-term ranges, we built T-LoCoH time use grids to estimate recursive movement by each individual. Comparing these grids to contemporaneous MODIS imagery, we investigated productivity's influence on short-term space use and recursion. Finally, we investigated patterns of recursion within a year's home range, measuring the time to return to the most intensively used patches.ResultsTwenty four-hour displacements at dawn were frequently smaller than 24-h displacements at dusk or at midday and midnight resting periods. Recursion analyses demonstrated that short-term recursion was most common in areas of median rather than maximum NDVI values. Investigated across a full year, recursion analysis showed rhinos most frequently returned to areas within 8-21 days, though visits were also seen separated by months likely suggesting seasonality in range use.ConclusionsOur results indicate that rhinos may frequently stay within the same area of their home ranges for days at a time, and possibly return to the same general area days in a row especially during morning foraging bouts. Recursion across larger time scales is also evident, and likely a contributing mechanism for maintaining open landscapes and browsing lawns of the savanna.

Neanderthals and modern humans both occupied the Levant for tens of thousands of years prior to the spread of modern humans into the rest of Eurasia and their replacement of the Neanderthals. That the inter-species boundary remained geographically localized for so long is a puzzle, particularly in light of the rapidity of its subsequent movement. Here, we propose that infectious-disease dynamics can explain the localization and persistence of the inter-species boundary. We further propose, and support with dynamical-systems models, that introgression-based transmission of alleles related to the immune system would have gradually diminished this barrier to pervasive inter-species interaction, leading to the eventual release of the inter-species boundary from its geographic localization. Asymmetries between the species in the characteristics of their associated 'pathogen packages' could have generated feedback that allowed modern humans to overcome disease burden earlier than Neanderthals, giving them an advantage in their subsequent spread into Eurasia.

Bacillus anthracis is a spore-forming, Gram-positive bacterium responsible for anthrax, an acute infection that most significantly affects grazing livestock and wild ungulates, but also poses a threat to human health. The geographic extent of B. anthracis is poorly understood, despite multi-decade research on anthrax epizootic and epidemic dynamics; many countries have limited or inadequate surveillance systems, even within known endemic regions. Here, we compile a global occurrence dataset of human, livestock and wildlife anthrax outbreaks. With these records, we use boosted regression trees to produce a map of the global distribution of B. anthracis as a proxy for anthrax risk. We estimate that 1.83 billion people (95% credible interval (CI): 0.59-4.16 billion) live within regions of anthrax risk, but most of that population faces little occupational exposure. More informatively, a global total of 63.8 million poor livestock keepers (95% CI: 17.5-168.6 million) and 1.1 billion livestock (95% CI: 0.4-2.3 billion) live within vulnerable regions. Human and livestock vulnerability are both concentrated in rural rainfed systems throughout arid and temperate land across Eurasia, Africa and North America. We conclude by mapping where anthrax risk could disrupt sensitive conservation efforts for wild ungulates that coincide with anthrax-prone landscapes.

Dynamic SEIR (Susceptible, Exposed, Infectious, Removed) compartmental models provide a tool for predicting the size and duration of both unfettered and managed outbreaks-the latter in the context of interventions such as case detection, patient isolation, vaccination and treatment. The reliability of this tool depends on the validity of key assumptions that include homogeneity of individuals and spatio-temporal homogeneity. Although the SEIR compartmental framework can easily be extended to include demographic (e.g. age) and additional disease (e.g. healthcare workers) classes, dependence of transmission rates on time, and metapopulation structure, fitting such extended models is hampered by both a proliferation of free parameters and insufficient or inappropriate data. This raises the question of how effective a tool the basic SEIR framework may actually be. We go some way here to answering this question in the context of the 2014-2015 outbreak of Ebola in West Africa by comparing fits of an SEIR time-dependent transmission model to both country- and district-level weekly incidence data. Our novel approach in estimating the effective-size-of-the-populations-at-risk ( Neff) and initial number of exposed individuals ( E0) at both district and country levels, as well as the transmission function parameters, including a time-to-halving-the-force-of-infection ( tf/2) parameter, provides new insights into this Ebola outbreak. It reveals that the estimate R0 ≈ 1.7 from country-level data appears to seriously underestimate R0 ≈ 3.3 - 4.3 obtained from more spatially homogeneous district-level data. Country-level data also overestimate tf/2 ≈ 22 weeks, compared with 8-10 weeks from district-level data. Additionally, estimates for the duration of individual infectiousness is around two weeks from spatially inhomogeneous country-level data compared with 2.4-4.5 weeks from spatially more homogeneous district-level data, which estimates are rather high compared with most values reported in the literature. This article is part of the theme issue 'Modelling infectious disease outbreaks in humans, animals and plants: approaches and important themes'. This issue is linked with the subsequent theme issue 'Modelling infectious disease outbreaks in humans, animals and plants: epidemic forecasting and control'.

The implementation of intelligent software to identify and classify objects and individuals in visual fields is a technology of growing importance to operatives in many fields, including wildlife conservation and management. To non-experts, the methods can be abstruse and the results mystifying. Here, in the context of applying cutting edge methods to classify wildlife species from camera-trap data, we shed light on the methods themselves and types of features these methods extract to make efficient identifications and reliable classifications. The current state of the art is to employ convolutional neural networks (CNN) encoded within deep-learning algorithms. We outline these methods and present results obtained in training a CNN to classify 20 African wildlife species with an overall accuracy of 87.5% from a dataset containing 111,467 images. We demonstrate the application of a gradient-weighted class-activation-mapping (Grad-CAM) procedure to extract the most salient pixels in the final convolution layer. We show that these pixels highlight features in particular images that in some cases are similar to those used to train humans to identify these species. Further, we used mutual information methods to identify the neurons in the final convolution layer that consistently respond most strongly across a set of images of one particular species. We then interpret the features in the image where the strongest responses occur, and present dataset biases that were revealed by these extracted features. We also used hierarchical clustering of feature vectors (i.e., the state of the final fully-connected layer in the CNN) associated with each image to produce a visual similarity dendrogram of identified species. Finally, we evaluated the relative unfamiliarity of images that were not part of the training set when these images were one of the 20 species "known" to our CNN in contrast to images of the species that were "unknown" to our CNN.

How a disease is transmitted affects our ability to determine R₀, the average number of new cases caused by an infectious host at the onset of an epidemic. R₀ becomes progressively more difficult to compute as transmission varies from directly transmitted diseases to diseases that are vector-borne to environmentally transmitted diseases. Pathogens responsible for diseases with environmental transmission are typically maintained in environmental reservoirs that exhibit a complex spatial distribution of local infectious zones (LIZs). Understanding host encounters with LIZs and pathogen persistence within LIZs is required for an accurate R₀ and modeling these contacts requires an integrated geospatial and dynamical systems approach. Here we review how interactions between host and pathogen populations and environmental reservoirs are driven by landscape-level variables, and synthesize the quantitative framework needed to formulate outbreak response and disease control.

Bovine tuberculosis (BTB) is endemic to the African buffalo (Syncerus caffer) of Hluhluwe-iMfolozi Park (HiP) and Kruger National Park, South Africa. In HiP, the disease has been actively managed since 1999 through a test-and-cull procedure targeting BTB-positive buffalo. Prior studies in Kruger showed associations between microsatellite alleles, BTB and body condition. A sex chromosomal meiotic drive, a form of natural gene drive, was hypothesized to be ultimately responsible. These associations indicate high-frequency occurrence of two types of male-deleterious alleles (or multiple-allele haplotypes). One type negatively affects body condition and BTB resistance in both sexes. The other type has sexually antagonistic effects: negative in males but positive in females. Here, we investigate whether a similar gene drive system is present in HiP buffalo, using 17 autosomal microsatellites and microsatellite-derived Y-chromosomal haplotypes from 401 individuals, culled in 2002-2004. We show that the association between autosomal microsatellite alleles and BTB susceptibility detected in Kruger, is also present in HiP. Further, Y-haplotype frequency dynamics indicated that a sex chromosomal meiotic drive also occurred in HiP. BTB was associated with negative selection of male-deleterious alleles in HiP, unlike positive selection in Kruger. Birth sex ratios were female-biased. We attribute negative selection and female-biased sex ratios in HiP to the absence of a Y-chromosomal sex-ratio distorter. This distorter has been hypothesized to contribute to positive selection of male-deleterious alleles and male-biased birth sex ratios in Kruger. As previously shown in Kruger, microsatellite alleles were only associated with male-deleterious effects in individuals born after wet pre-birth years; a phenomenon attributed to epigenetic modification. We identified two additional allele types: male-specific deleterious and beneficial alleles, with no discernible effect on females. Finally, we discuss how our findings may be used for breeding disease-free buffalo and implementing BTB test-and-cull programs.

ABSTRACT Animal movement paths are represented by point-location time series called relocation data. How well such paths can be simulated, when the rules governing movement depend on the internal state of individuals and environmental factors (both local and, when memory is involved, global) remains an open question. To answer this, we formulate and test models able to capture the essential statistics of multiphase versions of such paths (viz., movement-phase-specific step-length and turning-angle means, variances, auto-correlation, and cross correlation values), as well as broad scale movement patterns. The latter may include patchy coverage of the landscape, as well as the existence of home-range boundaries and gravitational centers-of-movement (e.g., centered around nests). Here we present a Numerus Model Builder implementation of two kinds of models: a high-frequency, multi-mode, biased, correlated random walk designed to simulate real movement data at a scale that permits simulation and identification of path segments that range from minutes to days; and a model that uses statistics extracted from relocation data—either empirical or simulated—to construct movement modes and phases at subhourly to daily scales. We evaluate how well our derived statistical movement model captures patterns produced by our more detailed simulation model as a way to evaluate how well derived statistical movement models may capture patterns occurring in empirical data.

Animal relocation data has recently become considerably more ubiquitous, finely structured (collection frequencies measured in minutes) and co-variate rich (physiology of individuals, environmental and landscape information, and accelerometer data). To better understand the impacts of ecological interactions, individual movement and disease on global change ecology, including wildlife management and conservation, it is important to have simulators that will provide demographic, movement, and epidemiology null models against which to compare patterns observed in empirical systems. Such models may then be used to develop quantitative narratives that enhance our intuition and understanding of the relationship between population structure and generative processes: in essence, along with empirical and experimental narratives, quantitative narratives are used to advance ecological epistemology. Here we describe a simulator that accounts for the influence of consumer-resource interactions, existence of social groups anchored around a central location, territoriality, group-switching behavior, and disease dynamics on population size. We use this simulator to develop new and reinforce existing quantitative narratives and point out areas for future study. Author summary The health and viability of species are of considerable concern to all nature lovers. Population models are central to our efforts to assess the numerical and ecological status of species and threats posed by climate change. Models, however, are crude caricatures of complex ecological systems. So how do we construct reliable assessment models able to capture processes essential to predicating the impacts of global change on population viability without getting tied up in their vast complexities? We broach this question and demonstrate how models focusing at the level of the individual (i.e., agent-based models) are tools for developing robust, narratives to augment narratives arising purely from empirical data sources and experimental outcomes. We do this in the context of nesting social groups, foraging for food, while exhibiting territoriality and group-switching behavior; and, we evaluate the impact of disease on the viability of such populations.

Background As GPS tags and data loggers have become lighter, cheaper, and longer-lasting, there has been a growing influx of data on animal movement. Simultaneously, methods of analyses and software to apply such methods to movement data have expanded dramatically. Even so, for many interdisciplinary researchers and managers without familiarity with the field of movement ecology and the open-source tools that have been developed, the analysis of movement data has remained an overwhelming challenge. Description Here we present stmove , an R package designed to take individual relocation data and generate a visually rich report containing a set of preliminary results that ecologists and managers can use to guide further exploration of their data. Not only does this package make report building and exploratory data analysis (EDA) simple for users who may not be familiar with the extent of available analytical tools, but it sets forth a framework of best practice analyses, which offers a common starting point for the interpretation of terrestrial movement data. Results Using data from African elephants ( Loxodonta africana ) collected in southern Africa, we demonstrate stmove ’s report building function through the main analyses included: path visualization, primary statistic calculation, summary in space and time, and space-use construction. Conclusions The stmove package provides consistency and increased accessibility to managers and researchers who are interested in movement analysis but who may be unfamiliar with the full scope of movement packages and analytical tools. If widely adopted, the package will promote comparability of results across movement ecology studies.

Epidemiological models are dominated by compartmental models, of which SIR formulations are the most commonly used. These formulations can be continuous or discrete (in either the state-variable values or time), deterministic or stochastic, or spatially homogeneous or heterogeneous, the latter often embracing a network formulation. Here we review the continuous and discrete deterministic and discrete stochastic formulations of the SIR dynamical systems models, and we outline how they can be easily and rapidly constructed using Numerus Model Builder, a graphically-driven coding platform. We also demonstrate how to extend these models to a metapopulation setting using NMB network and mapping tools.

Erlang differential equation models of epidemic processes provide more realistic disease-class transition dynamics from susceptible (S) to exposed (E) to infectious (I) and removed (R) categories than the ubiquitous SEIR model. The latter is itself is at one end of the spectrum of Erlang SE[Formula: see text]I[Formula: see text]R models with [Formula: see text] concatenated E compartments and [Formula: see text] concatenated I compartments. Discrete-time models, however, are computationally much simpler to simulate and fit to epidemic outbreak data than continuous-time differential equations, and are also much more readily extended to include demographic and other types of stochasticity. Here we formulate discrete-time deterministic analogs of the Erlang models, and their stochastic extension, based on a time-to-go distributional principle. Depending on which distributions are used (e.g. discretized Erlang, Gamma, Beta, or Uniform distributions), we demonstrate that our formulation represents both a discretization of Erlang epidemic models and generalizations thereof. We consider the challenges of fitting SE[Formula: see text]I[Formula: see text]R models and our discrete-time analog to data (the recent outbreak of Ebola in Liberia). We demonstrate that the latter performs much better than the former; although confining fits to strict SEIR formulations reduces the numerical challenges, but sacrifices best-fit likelihood scores by at least 7%.

The growing field of movement ecology uses high resolution movement data to analyze animal behavior across multiple scales: from individual foraging decisions to population-level space-use patterns. These analyses contribute to various subfields of ecology-inter alia behavioral, disease, landscape, resource, and wildlife-and facilitate facilitate novel exploration in fields ranging from conservation planning to public health. Despite the growing availability and general accessibility of animal movement data, much potential remains for the analytical methods of movement ecology to be incorporated in all types of geographic analyses. This review provides for the Geographical Information Sciences (GIS) community an overview of the most common movement metrics and methods of analysis employed by animal ecologists. Through illustrative applications, we emphasize the potential for movement analyses to promote transdisciplinary GIS/wildlife-ecology research.

Environmentally transmitted diseases are comparatively poorly understood and managed, and their ecology is particularly understudied. Here we identify challenges of studying environmental transmission and persistence with a six-sided interdisciplinary review of the biology of anthrax (Bacillus anthracis). Anthrax is a zoonotic disease capable of maintaining infectious spore banks in soil for decades (or even potentially centuries), and the mechanisms of its environmental persistence have been the topic of significant research and controversy. Where anthrax is endemic, it plays an important ecological role, shaping the dynamics of entire herbivore communities. The complex eco-epidemiology of anthrax, and the mysterious biology of Bacillus anthracis during its environmental stage, have necessitated an interdisciplinary approach to pathogen research. Here, we illustrate different disciplinary perspectives through key advances made by researchers working in Etosha National Park, a long-term ecological research site in Namibia that has exemplified the complexities of the enzootic process of anthrax over decades of surveillance. In Etosha, the role of scavengers and alternative routes (waterborne transmission and flies) has proved unimportant relative to the long-term persistence of anthrax spores in soil and their infection of herbivore hosts. Carcass deposition facilitates green-ups of vegetation to attract herbivores, potentially facilitated by the role of anthrax spores in the rhizosphere. The underlying seasonal pattern of vegetation, and herbivores' immune and behavioural responses to anthrax risk, interact to produce regular 'anthrax seasons' that appear to be a stable feature of the Etosha ecosystem. Through the lens of microbiologists, geneticists, immunologists, ecologists, epidemiologists, and clinicians, we discuss how anthrax dynamics are shaped at the smallest scale by population genetics and interactions within the bacterial communities up to the broadest scales of ecosystem structure. We illustrate the benefits and challenges of this interdisciplinary approach to disease ecology, and suggest ways anthrax might offer insights into the biology of other important pathogens. Bacillus anthracis, and the more recently emerged Bacillus cereus biovar anthracis, share key features with other environmentally transmitted pathogens, including several zoonoses and panzootics of special interest for global health and conservation efforts. Understanding the dynamics of anthrax, and developing interdisciplinary research programs that explore environmental persistence, is a critical step forward for understanding these emerging threats.

Context: Habitat selection is a fundamental process that shapes animal spatial ecology. Species with wide geographic distribution that occupy diverse habitats have to adapt their resource acquisition strategies to maximise their effectiveness under local ecological constraints, leading to intraspecific behavioural variability. Identifying environmental determinants of habitat use pattern and regional intraspecific differences advances our understanding of the ecological underpinnings of animal behaviour and is important in strategising effective conservation and management of free-ranging populations. Aims: The aim of this study was to assess individual heterogeneity of habitat selection and use by African elephants under different seasonal and ecological constraints, in order to better understand the processes underlying their spatial behaviour. Methods: We investigated the habitat selection pattern of 19 African elephants equipped with satellite-linked GPS-collars in two different ecosystems, resource-rich bushveld bordering Kruger National Park, South Africa (six individuals) and arid savannah of Etosha National Park, Namibia (13 individuals). By constructing individual-specific and population-level resource selection functions (RSFs), we examined seasonal differences of elephant habitat use pattern to identify the underlying ecological mechanisms. Key results: Elephants were attracted to surface water in both study areas; but when water availability decreased in arid environment, they showed individual-specific preference in using natural vs artificial water sources. Road networks enabled efficient travel among resource patches, but its use differed between individuals. Areas with higher and more predictable vegetation productivity were generally preferred by elephants in dry season, but in more competitive arid savannah system there were individual/group-specific seasonal differences in resource selection patterns, likely reflecting the social dynamics among individuals. At population-level, the habitat selection pattern was less apparent due to considerable intra-population variability. Conclusions: The substantial differences in model coefficients within and between our study populations demonstrate the spatio-behavioural plasticity of elephants under various environmental conditions and suggest that population-level RSFs may over-simplify elephants' socio-ecological complexity. Implications: Within the resource competition paradigm, individual-specific habitat selection may be as essential in maintaining population resilience as is the population-level pattern of resource use. Consequently, spatio-behavioural heterogeneity within and between populations should be accounted for in resource selection studies and all subsequent conservation management policies. Habitat use pattern of animals is shaped by their biological needs and ecological dynamics of their environment. We examined habitat selection of African elephants in two different ecosystems in southern Africa, and found that individual responses to seasonally fluctuating resources varied considerably within and between populations. We conclude that individual-specific habitat selection pattern is as essential in maintaining population resilience as is the population-level pattern, and the intra- and inter-population spatio-behavioural heterogeneity have to be accounted for in conservation management. Photograph by Scott Y. S. Chui. [ABSTRACT FROM AUTHOR]

Though epidemiology dates back to the 1700s, most mathematical representations of epidemics still use transmission rates averaged at the population scale, especially for wildlife diseases. In simplifying the contact process, we ignore the heterogeneities in host movements that complicate the real world, and overlook their impact on spatiotemporal patterns of disease burden. Movement ecology offers a set of tools that help unpack the transmission process, letting researchers more accurately model how animals within a population interact and spread pathogens. Analytical techniques from this growing field can also help expose the reverse process: how infection impacts movement behaviours, and therefore other ecological processes like feeding, reproduction, and dispersal. Here, we synthesise the contributions of movement ecology in disease research, with a particular focus on studies that have successfully used movement-based methods to quantify individual heterogeneity in exposure and transmission risk. Throughout, we highlight the rapid growth of both disease and movement ecology and comment on promising but unexplored avenues for research at their overlap. Ultimately, we suggest, including movement empowers ecologists to pose new questions, expanding our understanding of host-pathogen dynamics and improving our predictive capacity for wildlife and even human diseases.

Critical evaluation of the adequacy of ecological models is urgently needed to enhance their utility in developing theory and enabling environmental managers and policymakers to make informed decisions. Poorly supported management can have detrimental, costly or irreversible impacts on the environment and society. Here, we examine common issues in ecological modelling and suggest criteria for improving modelling frameworks. An appropriate level of process description is crucial to constructing the best possible model, given the available data and understanding of ecological structures. Model details unsupported by data typically lead to over parameterisation and poor model performance. Conversely, a lack of mechanistic details may limit a model's ability to predict ecological systems' responses to management. Ecological studies that employ models should follow a set of model adequacy assessment protocols that include: asking a series of critical questions regarding state and control variable selection, the determinacy of data, and the sensitivity and validity of analyses. We also need to improve model elaboration, refinement and coarse graining procedures to better understand the relevancy and adequacy of our models and the role they play in advancing theory, improving hind and forecasting, and enabling problem solving and management.

BackgroundContinued exploration of the performance of the recently proposed cross-validation-based approach for delimiting home ranges using the Time Local Convex Hull (T-LoCoH) method has revealed a number of issues with the original formulation.Main textHere we replace the ad hoc cross-validation score with a new formulation based on the total log probability of out-of-sample predictions. To obtain these probabilities, we interpret the normalized LoCoH hulls as a probability density. The application of the approach described here results in optimal parameter sets that differ dramatically from those selected using the original formulation. The derived metrics of home range size, mean revisitation rate, and mean duration of visit are also altered using the corrected formulation.ConclusionDespite these differences, we encourage the use of the cross-validation-based approach, as it provides a unifying framework governed by the statistical properties of the home ranges rather than subjective selections by the user.

In the African buffalo (Syncerus caffer) population of the Kruger National Park (South Africa) a primary sex-ratio distorter and a primary sex-ratio suppressor have been shown to occur on the Y chromosome. A subsequent autosomal microsatellite study indicated that two types of deleterious alleles with a negative effect on male body condition, but a positive effect on relative fitness when averaged across sexes and generations, occur genome-wide and at high frequencies in the same population. One type negatively affects body condition of both sexes, while the other acts antagonistically: it negatively affects male but positively affects female body condition. Here we show that high frequencies of male-deleterious alleles are attributable to Y-chromosomal distorter-suppressor pair activity and that these alleles are suppressed in individuals born after three dry pre-birth years, likely through epigenetic modification. Epigenetic suppression was indicated by statistical interactions between pre-birth rainfall, a proxy for parental body condition, and the phenotypic effect of homozygosity/heterozygosity status of microsatellites linked to male-deleterious alleles, while a role for the Y-chromosomal distorter-suppressor pair was indicated by between-sex genetic differences among pre-dispersal calves. We argue that suppression of male-deleterious alleles results in negative frequency-dependent selection of the Y distorter and suppressor; a prerequisite for a stable polymorphism of the Y distorter-suppressor pair. The Y distorter seems to be responsible for positive selection of male-deleterious alleles during resource-rich periods and the Y suppressor for positive selection of these alleles during resource-poor periods. Male-deleterious alleles were also associated with susceptibility to bovine tuberculosis, indicating that Kruger buffalo are sensitive to stressors such as diseases and droughts. We anticipate that future genetic studies on African buffalo will provide important new insights into gene fitness and epigenetic modification in the context of sex-ratio distortion and infectious disease dynamics.

[This corrects the article DOI: 10.1186/s40462-017-0110-4.].

BackgroundWith decreasing costs of GPS telemetry devices, data repositories of animal movement paths are increasing almost exponentially in size. A series of complex statistical tools have been developed in conjunction with this increase in data. Each of these methods offers certain improvements over previously proposed methods, but each has certain assumptions or shortcomings that make its general application difficult. In the case of the recently developed Time Local Convex Hull (T-LoCoH) method, the subjectivity in parameter selection serves as one of the primary impediments to its more widespread use. While there are certain advantages to the flexibility it offers for question-driven research, the lack of an objective approach for parameter selection may prevent some users from exploring the benefits of the method.MethodsHere we present a cross-validation-based approach for selecting parameter values to optimize the T-LoCoH algorithm. We demonstrate the utility of the approach using a case study from the Etosha National Park anthrax system.ResultsUtilizing the proposed algorithm, rather than the guidelines in the T-LoCoH documentation, results in significantly different values for derived site fidelity metrics.ConclusionsDue to its basis in principles of cross-validation, the application of this method offers a more objective approach than the relatively subjective guidelines set forth in the T-LoCoH documentation and enables a more accurate basis for the comparison of home ranges among individuals and species, as well as among studies.

Climate change is a well-documented driver of both wildlife extinction and disease emergence, but the negative impacts of climate change on parasite diversity are undocumented. We compiled the most comprehensive spatially explicit data set available for parasites, projected range shifts in a changing climate, and estimated extinction rates for eight major parasite clades. On the basis of 53,133 occurrences capturing the geographic ranges of 457 parasite species, conservative model projections suggest that 5 to 10% of these species are committed to extinction by 2070 from climate-driven habitat loss alone. We find no evidence that parasites with zoonotic potential have a significantly higher potential to gain range in a changing climate, but we do find that ectoparasites (especially ticks) fare disproportionately worse than endoparasites. Accounting for host-driven coextinctions, models predict that up to 30% of parasitic worms are committed to extinction, driven by a combination of direct and indirect pressures. Despite high local extinction rates, parasite richness could still increase by an order of magnitude in some places, because species successfully tracking climate change invade temperate ecosystems and replace native species with unpredictable ecological consequences.

African savanna vegetation is subject to extensive degradation as a result of rapid climate and land use change. To better understand these changes detailed assessment of vegetation structure is needed across an extensive spatial scale and at a fine temporal resolution. Applying remote sensing techniques to savanna vegetation is challenging due to sparse cover, high background soil signal, and difficulty to differentiate between spectral signals of bare soil and dry vegetation. In this paper, we attempt to resolve these challenges by analyzing time series of four MODIS Vegetation Products (VPs): Normalized Difference Vegetation Index (NDVI), Enhanced Vegetation Index (EVI), Leaf Area Index (LAI), and Fraction of Photosynthetically Active Radiation (FPAR) for Etosha National Park, a semiarid savanna in north-central Namibia. We create models to predict the density, cover, and biomass of the main savanna vegetation forms: grass, shrubs, and trees. To calibrate remote sensing data we developed an extensive and relatively rapid field methodology and measured herbaceous and woody vegetation during both the dry and wet seasons. We compared the efficacy of the four MODIS-derived VPs in predicting vegetation field measured variables. We then compared the optimal time span of VP time series to predict ground-measured vegetation. We found that Multiyear Partial Least Square Regression (PLSR) models were superior to single year or single date models. Our results show that NDVI-based PLSR models yield robust prediction of tree density (R2 =0.79, relative Root Mean Square Error, rRMSE=1.9%) and tree cover (R2 =0.78, rRMSE=0.3%). EVI provided the best model for shrub density (R2 =0.82) and shrub cover (R2 =0.83), but was only marginally superior over models based on other VPs. FPAR was the best predictor of vegetation biomass of trees (R2 =0.76), shrubs (R2 =0.83), and grass (R2 =0.91). Finally, we addressed an enduring challenge in the remote sensing of semiarid vegetation by examining the transferability of predictive models through space and time. Our results show that models created in the wetter part of Etosha could accurately predict trees' and shrubs' variables in the drier part of the reserve and vice versa. Moreover, our results demonstrate that models created for vegetation variables in the dry season of 2011 could be successfully applied to predict vegetation in the wet season of 2012. We conclude that extensive field data combined with multiyear time series of MODIS vegetation products can produce robust predictive models for multiple vegetation forms in the African savanna. These methods advance the monitoring of savanna vegetation dynamics and contribute to improved management and conservation of these valuable ecosystems.

Ecologists have traditionally focused on herbivore carcasses as study models in scavenging research. However, some observations of scavengers avoiding feeding on carnivore carrion suggest that different types of carrion may lead to differential pressures. Untested assumptions about carrion produced at different trophic levels could therefore lead ecologists to overlook important evolutionary processes and their ecological consequences. Our general goal was to investigate the use of mammalian carnivore carrion by vertebrate scavengers. In particular, we aimed to test the hypothesis that carnivore carcasses are avoided by other carnivores, especially at the intraspecific level, most likely to reduce exposure to parasitism. We take a three-pronged approach to study this principle by: (i) providing data from field experiments, (ii) carrying out evolutionary simulations of carnivore scavenging strategies under risks of parasitic infection, and (iii) conducting a literature-review to test two predictions regarding parasite life-history strategies. First, our field experiments showed that the mean number of species observed feeding at carcasses and the percentage of consumed carrion biomass were substantially higher at herbivore carcasses than at carnivore carcasses. This occurred even though the number of scavenger species visiting carcasses and the time needed by scavengers to detect carcasses were similar between both types of carcasses. In addition, we did not observe cannibalism. Second, our evolutionary simulations demonstrated that a risk of parasite transmission leads to the evolution of scavengers with generally low cannibalistic tendencies, and that the emergence of cannibalism-avoidance behaviour depends strongly on assumptions about parasite-based mortality rates. Third, our literature review indicated that parasite species potentially able to follow a carnivore-carnivore indirect cycle, as well as those transmitted via meat consumption, are rare in our study system. Our findings support the existence of a novel coevolutionary relation between carnivores and their parasites, and suggest that carnivore and herbivore carcasses play very different roles in food webs and ecosystems.