Decision rules for determining terrestrial movement and the consequences for filtering high-resolution global positioning system tracks: a case study using the African lion (Panthera leo).

The combined use of global positioning system (GPS) technology and motion sensors within the discipline of movement ecology has increased over recent years. This is particularly the case for instrumented wildlife, with many studies now opting to record parameters at high (infra-second) sampling frequencies. However, the detail with which GPS loggers can elucidate fine-scale movement depends on the precision and accuracy of fixes, with accuracy being affected by signal reception. We hypothesized that animal behaviour was the main factor affecting fix inaccuracy, with inherent GPS positional noise (jitter) being most apparent during GPS fixes for non-moving locations, thereby producing disproportionate error during rest periods. A movement-verified filtering (MVF) protocol was constructed to compare GPS-derived speed data with dynamic body acceleration, to provide a computationally quick method for identifying genuine travelling movement. This method was tested on 11 free-ranging lions (Panthera leo) fitted with collar-mounted GPS units and tri-axial motion sensors recording at 1 and 40 Hz, respectively. The findings support the hypothesis and show that distance moved estimates were, on average, overestimated by greater than 80% prior to GPS screening. We present the conceptual and mathematical protocols for screening fix inaccuracy within high-resolution GPS datasets and demonstrate the importance that MVF has for avoiding inaccurate and biased estimates of movement.

How often should dead-reckoned animal movement paths be corrected for drift?

BACKGROUND: Understanding what animals do in time and space is important for a range of ecological questions, however accurate estimates of how animals use space is challenging. Within the use of animal-attached tags, radio telemetry (including the Global Positioning System, 'GPS') is typically used to verify an animal's location periodically. Straight lines are typically drawn between these 'Verified Positions' ('VPs') so the interpolation of space-use is limited by the temporal and spatial resolution of the system's measurement. As such, parameters such as route-taken and distance travelled can be poorly represented when using VP systems alone. Dead-reckoning has been suggested as a technique to improve the accuracy and resolution of reconstructed movement paths, whilst maximising battery life of VP systems. This typically involves deriving travel vectors from motion sensor systems and periodically correcting path dimensions for drift with simultaneously deployed VP systems. How often paths should be corrected for drift, however, has remained unclear. METHODS AND RESULTS: Here, we review the utility of dead-reckoning across four contrasting model species using different forms of locomotion (the African lion Panthera leo, the red-tailed tropicbird Phaethon rubricauda, the Magellanic penguin Spheniscus magellanicus, and the imperial cormorant Leucocarbo atriceps). Simulations were performed to examine the extent of dead-reckoning error, relative to VPs, as a function of Verified Position correction (VP correction) rate and the effect of this on estimates of distance moved. Dead-reckoning error was greatest for animals travelling within air and water. We demonstrate how sources of measurement error can arise within VP-corrected dead-reckoned tracks and propose advancements to this procedure to maximise dead-reckoning accuracy. CONCLUSIONS: We review the utility of VP-corrected dead-reckoning according to movement type and consider a range of ecological questions that would benefit from dead-reckoning, primarily concerning animal-barrier interactions and foraging strategies.

The future of sustainability in the context of COVID-19.

The COVID-19 pandemic is a global crisis emanating both from a virus (SARS-CoV-2) and from the drastic actions to contain it. Here, we reflect on the immediate responses of most world powers amid the pandemic chaos: totalitarian surveillance and nationalist isolation. Drawing on published literature, we consider measures such as wildlife-use bans, lockdowns and travel restrictions, along with their reverberations for people, economies and the planet. Our synthesis highlights significant shortfalls of applying command-and-control tactics in emergencies. For one, heavy-handed bans risk enormous unintended consequences and tend to fail if they lack legitimacy or clash with people's values. Furthermore, reactive and myopic strategies typically view the pandemic as a stand-alone crisis, rather than unravelling the complex interplay of nature-society interactions through which zoonotic diseases originate. A return to adaptive management approaches that recognise root causes and foster socio-ecological resilience will be essential to improve human and planetary health and mitigate future pandemics.

Ecological drivers of female lion (Panthera leo) reproduction in the Kruger National Park.

The role of social cues in the reproduction of social mammals, particularly carnivores, has been thoroughly studied and documented in literature. However, environmental cues such as resources of water, food, and shelter have been identified to a lesser extent. Pregnant lions (Panthera leo) are notoriously secretive during the final stages of pregnancy and postpartum. Behavioral indicators depicted by movement patterns obtained by remote detection of collared female lions in the Kruger National Park were necessary for the monitoring of birth timing. Over the study period, eight plus a potential three parturition incidences of collared females were recorded. Of the variables measured (step length, range size, duration, prey biomass, and rainfall), range size during the month of parturition was the most indicative movement pattern of a successful birth. By backdating the potential birth month of the litters, date of conception was calculated and our results revealed a correlation between the birthing peaks of preferred prey during the month of conception. Birth timing in conjunction with remote sensing and ecological factors were thus identified behaviors associated with denning.

Age structure changes indicate direct and indirect population impacts in illegally harvested black rhino.

Overharvesting affects the size and growth of wildlife populations and can impact population trajectories. Overharvesting can also severely alter population structure and may result in changes in spatial organisation, social dynamics and recruitment. Understanding the relationship between overharvesting and population growth is therefore crucial for the recovery of exploited species. The black rhinoceros (Diceros bicornis; black rhino) is a long-lived megaherbivore native to sub-Saharan Africa, listed as Critically Endangered on the IUCN Red List of Threatened Species. Since 2009, the targeted illegal killing of rhino for their horns has escalated dramatically in South Africa. Given their slow life trajectories, spatial structure and social dynamics, black rhino may be susceptible to both direct and indirect impacts of overharvesting. Our study compared black rhino demography before and during extensive poaching to understand the impact of illegal killing. The population exhibited significant changes in age structure after four years of heavy poaching; these changes were primarily explained by a decrease in the proportion of calves over time. Population projections incorporating both direct poaching removals and decreased fecundity/recruitment were most similar to the observed demographic profile in 2018, suggesting that indirect impacts are also contributing to the observed population trajectory. These indirect impacts are likely a result of decreased density, through processes such as reduced mate-finding, population disturbance and/or increased calf predation. This study illustrates the combined effect of direct and indirect impacts on an endangered species, providing a more comprehensive approach by which to evaluate exploited populations.

Top-down and bottom-up processes to implement biological monitoring in protected areas.

Achieving biodiversity conservation objectives and targets requires environmental management agencies to undertake monitoring. Several management practices have been used but are they successful? Using SANParks as an example, we focus on three cases of national parks that use different techniques to implement environmental monitoring. We assessed a top-down, bottom-up and an integrated approach to setting targets using Table Mountain, Mokala and Bontebok National Parks respectively. Attainment of national and international objectives from State of Biodiversity assessment scores and achievement of objectives within individual Park Management Plans served as measurement criteria. We highlight several reasons for lack of achievement of objectives and targets for protected areas and illustrate that setting of objectives and monitoring must have complete full integration into park operations, both on staff and budget scales, rather than be seen as an external function from service divisions within parks or external service providers. An integrated approach to setting of objectives and targets of national and international importance is best achieved via the robust implementation of strategic adaptive management with interventions, measurement, reflection and adaptation integrated as learning by doing. Our results may not be unique but only a few organizations implement adaptive management in its full context.

Species-specific drought impacts on black and white rhinoceroses.

Unrelenting poaching to feed the illegal trafficking of rhinoceros (rhino) horn remains the principle threat to the persistence of south-central black and southern white rhino that live in the Kruger National Park (Kruger), South Africa. Other global environmental change drivers, such as unpredictable climatic conditions, impose additional uncertainties on the management and persistence of these species. The drought experienced in Kruger over the 2015/2016 rainy season may have affected rhino population growth and thus added an additional population pressure to the poaching pressure already occurring. Under drought conditions, reduced grass biomass predicts increased natural deaths and a subsequent decrease in birth rate for the grazing white rhino. Such variance in natural death and birth rates for the browsing black rhino are not expected under these conditions. We evaluated these predictions using rhino population survey data from 2013 to 2017. Comparisons of natural deaths and birth rates between pre- (2013/2014 and 2014/15), during- (2015/2016) and post-drought (2016/2017) periods in Kruger showed increased natural mortality and decreased births for white rhino, but no significant changes for black rhino, supporting our predictions. As a result, despite reduced poaching rates, the total mortality rate of white rhino remains significantly higher than the birth rate. Decreased poaching, decreased natural deaths and no apparent drought effects in black rhino resulted in a lower total mortality rate than the estimated birth rate in 2017. Active biological management and traditional anti-poaching initiatives together therefore represent the most likely way to buffer the impacts of decreased population growth through climate change and wildlife crime on the persistence of rhinos.

Climate change leads to increasing population density and impacts of a key island invader.

The considerable threats of invasive rodents to island biodiversity are likely to be compounded by climate change. Forecasts for such interactions have been most pronounced for the Southern Ocean islands where ameliorating conditions are expected to decrease thermal and resource restrictions on rodents. Firm evidence for changing rodent populations in response to climate change, and demonstrations of associated impacts on the terrestrial environment, are nonetheless entirely absent for the region. Using data collected over three decades on sub-Antarctic Marion Island, we tested empirically whether mouse populations have changed through time and whether these changes can be associated significantly with changing abiotic conditions. Changes in invertebrate populations, which have previously been attributed to mouse predation, but with little explicit demographic analysis, were also examined to determine whether they can be associated with changing mouse populations. The total number of mice on the island at annual peak density increased by 430.0% between 1979-1980 and 2008-2011. This increase was due to an advanced breeding season, which was robustly related to the number of precipitation-free days during the non-breeding season. Mice directly reduced invertebrate densities, with biomass losses of up to two orders of magnitude in some habitats. Such invertebrate declines are expected to have significant consequences for ecosystem processes over the long term. Our results demonstrate that as climate change continues to create ameliorating conditions for invasive rodents on sub-Antarctic islands, the severity of their impacts will increase. They also emphasize the importance of rodent eradication for the restoration of invaded islands.

Finding politically feasible conservation policies: the case of wildlife trafficking.

Conservation management is of increasing importance in ecology as most ecosystems nowadays are essentially managed ecosystems. Conservation managers work within a political-ecological system when they develop and attempt to implement a conservation plan that is designed to meet particular conservation goals. In this article, we develop a decision support tool that can identify a conservation policy for a managed wildlife population that is both sustainable and politically feasible. Part of our tool consists of a simulation model composed of interacting influence diagrams. We build, fit, and use our tool on the case of rhino horn trafficking between South Africa and Asia. Using these diagrams, we show how a rhino poacher's belief system can be modified by such a policy and locate it in a perceived risks-benefits space before and after policy implementation. We statistically fit our model to observations on group actions and rhino abundance. We then use this fitted model to compute a politically feasible conservation policy.

Simplified large African carnivore density estimators from track indices.

BACKGROUND: The range, population size and trend of large carnivores are important parameters to assess their status globally and to plan conservation strategies. One can use linear models to assess population size and trends of large carnivores from track-based surveys on suitable substrates. The conventional approach of a linear model with intercept may not intercept at zero, but may fit the data better than linear model through the origin. We assess whether a linear regression through the origin is more appropriate than a linear regression with intercept to model large African carnivore densities and track indices. METHODS: We did simple linear regression with intercept analysis and simple linear regression through the origin and used the confidence interval for ß in the linear model y = αx + ß, Standard Error of Estimate, Mean Squares Residual and Akaike Information Criteria to evaluate the models. RESULTS: The Lion on Clay and Low Density on Sand models with intercept were not significant (P > 0.05). The other four models with intercept and the six models thorough origin were all significant (P < 0.05). The models using linear regression with intercept all included zero in the confidence interval for ß and the null hypothesis that ß = 0 could not be rejected. All models showed that the linear model through the origin provided a better fit than the linear model with intercept, as indicated by the Standard Error of Estimate and Mean Square Residuals. Akaike Information Criteria showed that linear models through the origin were better and that none of the linear models with intercept had substantial support. DISCUSSION: Our results showed that linear regression through the origin is justified over the more typical linear regression with intercept for all models we tested. A general model can be used to estimate large carnivore densities from track densities across species and study areas. The formula observed track density = 3.26 × carnivore density can be used to estimate densities of large African carnivores using track counts on sandy substrates in areas where carnivore densities are 0.27 carnivores/100 km2 or higher. To improve the current models, we need independent data to validate the models and data to test for non-linear relationship between track indices and true density at low densities.

Combating Rhino Horn Trafficking: The Need to Disrupt Criminal Networks.

The onslaught on the World's wildlife continues despite numerous initiatives aimed at curbing it. We build a model that integrates rhino horn trade with rhino population dynamics in order to evaluate the impact of various management policies on rhino sustainability. In our model, an agent-based sub-model of horn trade from the poaching event up through a purchase of rhino horn in Asia impacts rhino abundance. A data-validated, individual-based sub-model of the rhino population of South Africa provides these abundance values. We evaluate policies that consist of different combinations of legal trade initiatives, demand reduction marketing campaigns, increased anti-poaching measures within protected areas, and transnational policing initiatives aimed at disrupting those criminal syndicates engaged in horn trafficking. Simulation runs of our model over the next 35 years produces a sustainable rhino population under only one management policy. This policy includes both a transnational policing effort aimed at dismantling those criminal networks engaged in rhino horn trafficking-coupled with increases in legal economic opportunities for people living next to protected areas where rhinos live. This multi-faceted approach should be the focus of the international debate on strategies to combat the current slaughter of rhino rather than the binary debate about whether rhino horn trade should be legalized. This approach to the evaluation of wildlife management policies may be useful to apply to other species threatened by wildlife trafficking.

Continent-wide survey reveals massive decline in African savannah elephants.

African elephants (Loxodonta africana) are imperiled by poaching and habitat loss. Despite global attention to the plight of elephants, their population sizes and trends are uncertain or unknown over much of Africa. To conserve this iconic species, conservationists need timely, accurate data on elephant populations. Here, we report the results of the Great Elephant Census (GEC), the first continent-wide, standardized survey of African savannah elephants. We also provide the first quantitative model of elephant population trends across Africa. We estimated a population of 352,271 savannah elephants on study sites in 18 countries, representing approximately 93% of all savannah elephants in those countries. Elephant populations in survey areas with historical data decreased by an estimated 144,000 from 2007 to 2014, and populations are currently shrinking by 8% per year continent-wide, primarily due to poaching. Though 84% of elephants occurred in protected areas, many protected areas had carcass ratios that indicated high levels of elephant mortality. Results of the GEC show the necessity of action to end the African elephants' downward trajectory by preventing poaching and protecting habitat.

Estimating wildlife disease dynamics in complex systems using an Approximate Bayesian Computation framework.

Emerging infectious diseases of wildlife are of increasing concern to managers and conservation policy makers, but are often difficult to study and predict due to the complexity of host-disease systems and a paucity of empirical data. We demonstrate the use of an Approximate Bayesian Computation statistical framework to reconstruct the disease dynamics of bovine tuberculosis in Kruger National Park's lion population, despite limited empirical data on the disease's effects in lions. The modeling results suggest that, while a large proportion of the lion population will become infected with bovine tuberculosis, lions are a spillover host and long disease latency is common. In the absence of future aggravating factors, bovine tuberculosis is projected to cause a lion population decline of ~3% over the next 50 years, with the population stabilizing at this new equilibrium. The Approximate Bayesian Computation framework is a new tool for wildlife managers. It allows emerging infectious diseases to be modeled in complex systems by incorporating disparate knowledge about host demographics, behavior, and heterogeneous disease transmission, while allowing inference of unknown system parameters.

Conservation Risks: When Will Rhinos be Extinct?

We develop a risk intelligence system for biodiversity enterprises. Such enterprises depend on a supply of endangered species for their revenue. Many of these enterprises, however, cannot purchase a supply of this resource and are largely unable to secure the resource against theft in the form of poaching. Because replacements are not available once a species becomes extinct, insurance products are not available to reduce the risk exposure of these enterprises to an extinction event. For many species, the dynamics of anthropogenic impacts driven by economic as well as noneconomic values of associated wildlife products along with their ecological stressors can help meaningfully predict extinction risks. We develop an agent/individual-based economic-ecological model that captures these effects and apply it to the case of South African rhinos. Our model uses observed rhino dynamics and poaching statistics. It seeks to predict rhino extinction under the present scenario. This scenario has no legal horn trade, but allows live African rhino trade and legal hunting. Present rhino populations are small and threatened by a rising onslaught of poaching. This present scenario and associated dynamics predicts continued decline in rhino population size with accelerated extinction risks of rhinos by 2036. Our model supports the computation of extinction risks at any future time point. This capability can be used to evaluate the effectiveness of proposed conservation strategies at reducing a species' extinction risk. Models used to compute risk predictions, however, need to be statistically estimated. We point out that statistically fitting such models to observations will involve massive numbers of observations on consumer behavior and time-stamped location observations on thousands of animals. Finally, we propose Big Data algorithms to perform such estimates and to interpret the fitted model's output.

Disruption of Rhino Demography by Poachers May Lead to Population Declines in Kruger National Park, South Africa.

The onslaught on the World's rhinoceroses continues despite numerous initiatives aimed at curbing it. When losses due to poaching exceed birth rates, declining rhino populations result. We used previously published estimates and growth rates for black rhinos (2008) and white rhinos (2010) together with known poaching trends at the time to predict population sizes and poaching rates in Kruger National Park, South Africa for 2013. Kruger is a stronghold for the south-eastern black rhino and southern white rhino. Counting rhinos on 878 blocks 3x3 km in size using helicopters, estimating availability bias and collating observer and detectability biases allowed estimates using the Jolly's estimator. The exponential escalation in number of rhinos poached per day appears to have slowed. The black rhino estimate of 414 individuals (95% confidence interval: 343-487) was lower than the predicted 835 individuals (95% CI: 754-956). The white rhino estimate of 8,968 individuals (95% CI: 8,394-9,564) overlapped with the predicted 9,417 individuals (95% CI: 7,698-11,183). Density- and rainfall-dependent responses in birth- and death rates of white rhinos provide opportunities to offset anticipated poaching effects through removals of rhinos from high density areas to increase birth and survival rates. Biological management of rhinos, however, need complimentary management of the poaching threat as present poaching trends predict detectable declines in white rhino abundances by 2018. Strategic responses such as anti-poaching that protect supply from illegal harvesting, reducing demand, and increasing supply commonly require crime network disruption as a first step complimented by providing options for alternative economies in areas abutting protected areas.

Evaluating the status of African wild dogs Lycaon pictus and cheetahs Acinonyx jubatus through tourist-based photographic surveys in the Kruger National Park [corrected].

The Kruger National Park is a stronghold for African wild dog Lycaon pictus and cheetah Acinonyx jubatus conservation in South Africa. Tourist photographic surveys have been used to evaluate the minimum number of wild dogs and cheetahs alive over the last two decades. Photographic-based capture-recapture techniques for open populations were used on data collected during a survey done in 2008/9. Models were run for the park as a whole and per region (northern, central, southern). A total of 412 (329-495; SE 41.95) cheetahs and 151 (144-157; SE 3.21) wild dogs occur in the Kruger National Park. Cheetah capture probabilities were affected by time (number of entries) and sex, whereas wild dog capture probabilities were affected by the region of the park. When plotting the number of new individuals identified against the number of entries received, the addition of new wild dogs to the survey reached an asymptote at 210 entries, but cheetahs did not reach an asymptote. The cheetah population of Kruger appears to be acceptable, while the wild dog population size and density are of concern. The effectiveness of tourist-based surveys for estimating population sizes through capture-recapture analyses is shown.

Effects of the number of people on efficient capture and sample collection: a lion case study.

Certain carnivore research projects and approaches depend on successful capture of individuals of interest. The number of people present at a capture site may determine success of a capture. In this study 36 lion capture cases in the Kruger National Park were used to evaluate whether the number of people present at a capture site influenced lion response rates and whether the number of people at a sampling site influenced the time it took to process the collected samples. The analyses suggest that when nine or fewer people were present, lions appeared faster at a call-up locality compared with when there were more than nine people. The number of people, however, did not influence the time it took to process the lions. It is proposed that efficient lion capturing should spatially separate capture and processing sites and minimise the number of people at a capture site.

Lack of spatial and behavioral responses to immunocontraception application in African elephants (Loxodonta africana).

Opinions are divided as to whether human intervention to control elephant (Loxodonta africana) population growth is desirable, partly because of elephant welfare concerns. Female contraception through immunization with porcine zona pellucida (PZP) proteins is viable. The effects of sustained use and application of the PZP vaccine on elephant behavioral and spatial responses were examined by evaluating herd ranging, fission-fusion dynamics, association patterns, and reproductive and sexual behaviors. Minimal change was anticipated as a result of long calf dependence on and association with cows, a reduced but not indefinite 0% growth rate and the known mechanism of action of PZP vaccines, and minimal expected change in resource requirements necessitating behavioral or spatial use adaptations. Although behavioral effects identified in previous hormonal contraceptive trials were evident, it was demonstrated that immunocontraception caused no prolonged behavioral, social, or spatial changes over the 11-yr study period. Individually identified elephants were monitored from 1999 to 2011. Minimal, short-term social disruption, with temporary changes to the herds' core ranges, was observed during the annual treatment events, particularly in the first three treatment years, when vaccinations were conducted exclusively from the ground. Thereafter, when vaccinations were conducted aerially, minor disruptions were confined to the morning of administration only. Despite sustained treatments resulting in demographic changes of fewer calves being born, treatments did not alter spatial range use, and no adverse interherd-intraherd relations were observed. Similarly, resource requirements did not change as calving still occurred, although in fewer numbers. It was concluded that PZP immunocontraception has no detectable behavioral or social consequences in elephants over the course of 11 yr, providing a convincing argument for the use of sustained immunocontraception in the medium to long term as an important tool for elephant management. Behavioral consequences of alternative management approaches should all receive similar scrutiny to enable managers to make informed decisions when weighing management interventions.

Anthropogenic influences on conservation values of white rhinoceros.

White rhinoceros (rhinos) is a keystone conservation species and also provides revenue for protection agencies. Restoring or mimicking the outcomes of impeded ecological processes allows reconciliation of biodiversity and financial objectives. We evaluate the consequences of white rhino management removal, and in recent times, poaching, on population persistence, regional conservation outcomes and opportunities for revenue generation. In Kruger National Park, white rhinos increased from 1998 to 2008. Since then the population may vary non-directionally. In 2010, we estimated 10,621 (95% CI: 8,767-12,682) white rhinos using three different population estimation methods. The desired management effect of a varying population was detectable after 2008. Age and sex structures in sink areas (focal rhino capture areas) were different from elsewhere. This comes from relatively more sub-adults being removed by managers than what the standing age distribution defined. Poachers in turn focused on more adults in 2011. Although the effect of poaching was not detectable at the population level given the confidence intervals of estimates, managers accommodated expected poaching annually and adapted management removals. The present poaching trend predicts that 432 white rhinos may be poached in Kruger during 2012. The white rhino management model mimicking outcomes of impeded ecological processes predicts 397 rhino management removals are required. At present poachers may be doing "management removals," but conservationists have no opportunity left to contribute to regional rhino conservation strategies or generate revenue through white rhino sales. In addition, continued trends in poaching predict detectable white rhino declines in Kruger National Park by 2016. Our results suggest that conservationists need innovative approaches that reduce financial incentives to curb the threats that poaching poses to several conservation values of natural resources such as white rhinos.