Craig J. Tambling

From moonlight to movement and synchronized randomness: Fourier and wavelet analyses of animal location time series data

High-resolution animal location data are increasingly available, requiring analytical approaches and statistical tools that can accommodate the temporal structure and transient dynamics (non-stationarity) inherent in natural systems. Traditional analyses often assume uncorrelated or weakly correlated temporal structure in the velocity (net displacement) time series constructed using sequential location data. We propose that frequency and time-frequency domain methods, embodied by Fourier and wavelet transforms, can serve as useful probes in early investigations of animal movement data, stimulating new ecological insight and questions. We introduce a novel movement model with time-varying parameters to study these methods in an animal movement context. Simulation studies show that the spectral signature given by these methods provides a useful approach for statistically detecting and characterizing temporal dependency in animal movement data. In addition, our simulations provide a connection between the spectral signatures observed in empirical data with null hypotheses about expected animal activity. Our analyses also show that there is not a specific one-to-one relationship between the spectral signatures and behavior type and that departures from the anticipated signatures are also informative. Box plots of net displacement arranged by time of day and conditioned on common spectral properties can help interpret the spectral signatures of empirical data. The first case study is based on the movement trajectory of a lion (Panthera leo) that shows several characteristic daily activity sequences, including an active-rest cycle that is correlated with moonlight brightness. A second example based on six pairs of African buffalo (Syncerus caffer) illustrates the use of wavelet coherency to show that their movements synchronize when they are within approximately 1 km of each other, even when individual movement was best described as an uncorrelated random walk, providing an important spatial baseline of movement synchrony and suggesting that local behavioral cues play a strong role in driving movement patterns. We conclude with a discussion about the role these methods may have in guiding appropriately flexible probabilistic models connecting movement with biotic and abiotic covariates.

Ecological and financial impacts of illegal bushmeat trade in Zimbabwe

TRAFFIC Southern and East Africa, the European Union, Wilderness Trust, Chicago Board of Trade and the supporters of the African Wildlife Conservation Fund.

Underperformance of African Protected Area Networks and the Case for New Conservation Models: Insights from Zambia

Many African protected areas (PAs) are not functioning effectively. We reviewed the performance of Zambia’s PA network and provide insights into how their effectiveness might be improved. Zambia’s PAs are under-performing in ecological, economic and social terms. Reasons include: a) rapidly expanding human populations, poverty and open-access systems in Game Management Areas (GMAs) resulting in widespread bushmeat poaching and habitat encroachment; b) underfunding of the Zambia Wildlife Authority (ZAWA) resulting in inadequate law enforcement; c) reliance of ZAWA on extracting revenues from GMAs to cover operational costs which has prevented proper devolution of user-rights over wildlife to communities; d) on-going marginalization of communities from legal benefits from wildlife; e) under-development of the photo-tourism industry with the effect that earnings are limited to a fraction of the PA network; f) unfavourable terms and corruption which discourage good practice and adequate investment by hunting operators in GMAs; g) blurred responsibilities regarding anti-poaching in GMAs resulting in under-investment by all stakeholders. The combined effect of these challenges has been a major reduction in wildlife densities in most PAs and the loss of habitat in GMAs. Wildlife fares better in areas with investment from the private and/or NGO sector and where human settlement is absent. There is a need for: elevated government funding for ZAWA; greater international donor investment in protected area management; a shift in the role of ZAWA such that they focus primarily on national parks while facilitating the development of wildlife-based land uses by other stakeholders elsewhere; and new models for the functioning of GMAs based on joint-ventures between communities and the private and/or NGO sector. Such joint-ventures should provide defined communities with ownership of land, user-rights over wildlife and aim to attract long-term private/donor investment. These recommendations are relevant for many of the under-funded PAs occurring in other African countries.

An objective approach to determining the weight ranges of prey preferred by and accessible to the five large African carnivores.

Broad-scale models describing predator prey preferences serve as useful departure points for understanding predator-prey interactions at finer scales. Previous analyses used a subjective approach to identify prey weight preferences of the five large African carnivores, hence their accuracy is questionable. This study uses a segmented model of prey weight versus prey preference to objectively quantify the prey weight preferences of the five large African carnivores. Based on simulations of known predator prey preference, for prey species sample sizes above 32 the segmented model approach detects up to four known changes in prey weight preference (represented by model break-points) with high rates of detection (75% to 100% of simulations, depending on number of break-points) and accuracy (within 1.3±4.0 to 2.7±4.4 of known break-point). When applied to the five large African carnivores, using carnivore diet information from across Africa, the model detected weight ranges of prey that are preferred, killed relative to their abundance, and avoided by each carnivore. Prey in the weight ranges preferred and killed relative to their abundance are together termed “accessible prey”. Accessible prey weight ranges were found to be 14–135 kg for cheetah Acinonyx jubatus, 1–45 kg for leopard Panthera pardus, 32–632 kg for lion Panthera leo, 15–1600 kg for spotted hyaena Crocuta crocuta and 10–289 kg for wild dog Lycaon pictus. An assessment of carnivore diets throughout Africa found these accessible prey weight ranges include 88±2% (cheetah), 82±3% (leopard), 81±2% (lion), 97±2% (spotted hyaena) and 96±2% (wild dog) of kills. These descriptions of prey weight preferences therefore contribute to our understanding of the diet spectrum of the five large African carnivores. Where datasets meet the minimum sample size requirements, the segmented model approach provides a means of determining, and comparing, the prey weight range preferences of any carnivore species.

Methods for Locating African Lion Kills Using Global Positioning System Movement Data

Abstract Knowledge of the range, behavior, and feeding habits of large carnivores is fundamental to their successful conservation. Traditionally, the best method to obtain feeding data is through continuous observation, which is not always feasible. Reliable automated methods are needed to obtain sample sizes sufficient for statistical inference. Identification of large carnivore kill sites using Global Positioning System (GPS) data is gaining popularity. We assessed performance of generalized linear regression models (GLM) versus classification trees (CT) in a multipredator, multiprey African savanna ecosystem. We applied GLMs and CTs to various combinations of distance-traveled data, cluster durations, and environmental factors to predict occurrence of 234 female African lion (Panthera leo) kill sites from 1,477 investigated GPS clusters. Ratio of distance moved 24 hours before versus 24 hours after a cluster was the most important predictor variable in both GLM and CT analysis. In all cases, GLMs outperformed our cost-complexity-pruned CTs in their discriminative ability to separate kill from nonkill sites. Generalized linear models provided a good framework for kill-site identification that incorporates a hierarchal ordering of cluster investigation and measures to assess trade-offs between classification accuracy and time constraints. Implementation of GLMs within an adaptive sampling framework can considerably increase efficiency of locating kill sites, providing a cost-effective method for increasing sample sizes of kill data.

Minimum prey and area requirements of the Vulnerable cheetah Acinonyx jubatus: implications for reintroduction and management of the species in South Africa

In South Africa there are efforts to manage reintroduced subpopulations of the Vulnerable cheetah Acinonyx jubatus in small reserves (10-1,000 km 2 )a s a managed metapopulation. We estimated areas required to support cheetahs given varying prey densities, prey profiles and presence/absence of competing predators. A recent population and habitat viability assessment in- dicated that 20 subpopulations of 10 cheetahs or 10 sub- populations of 15 cheetahs are required to retain 90% of the heterozygosity of free-ranging cheetahs and to overcome stochastic events in the absence or presence of lions Panthera leo, respectively. We estimate that 203 - SE 42 km 2 (range 48-466 km 2 ) is required to support 10 cheetahs in the absence of lions, whereas 703 - SE 311 km 2 (166-2,806 km 2 ) is required to support 15 cheetahs given equal numbers of lions, and 2,424 - SE 890 km 2 (727-3,739 km 2 ) given equal numbers of leopards Panthera pardus, spotted hyaenas Crocuta crocuta, wild dogs Lycaon pictus and lions. Existing subpopulations of cheetahs generally occur at densities higher than our mean predicted densities but usually within the range of predicted densities. The large area requirements of cheetahs have implications for the development of the managed metapopulation. Sourcing reintroduction sites of the sizes required to support recommended subpopulation sizes will be difficult. Consequently, innovative measures to increase the carrying capacity of reserves for cheetahs and/or to enlarge reserves will be required. Managers may be forced to stock cheetahs close to or beyond the carrying capacity of their reserves. Consequently, careful management of rein- troduced subpopulations will be required to prevent declines in prey populations.

Habitat selection by large herbivores in a southern African savanna: the relative roles of bottom‐up and top‐down forces

This research was supported by the National Science Foundation of the United States (DEB 0516094 to A. Knapp and DEB 0516145 to M. Smith). Support from the James S. McDonnell Foundation and a United States National Science Foundation Grant (DEB 0090323) to W. Getz at the University of California, Berkeley funded the collars and field component on lion predation. D. Burkepile and C. Burns were supported, in part, by the Brown Postdoctoral Research Fellowship at Yale University.

The impact of power line-related mortality on the Cape Vulture Gyps coprotheres in a part of its range, with an emphasis on electrocution

The global population of the Cape Vulture Gyps coprotheres , a threatened southern African endemic, is known to be impacted by electrocutions and collisions on power line infrastructure, but to date this impact has not been estimated or quantified. Using data in a national database from the period prior to our study, conducted in the Eastern Cape Province of South Africa, we estimated a mean annual mortality rate from power line-related mortality of around 14 vultures per year. After applying an adjusted rate based on the results of a landowner survey, this estimate increased to around 80 vultures per year (i.e. a 5.7 fold increase). For a number of reasons, the estimated mean annual mortality rate is considered to under-represent the true situation, and must therefore be considered a minimum value. A simple model was constructed and run to investigate the potential impact of the mortality rate from electrocution on the study population. It distinguishes between vulture subpopulations in areas of high and low electrocution threat, and a migratory subpopulation that moves between these two areas. The model, simulated over 50 years and applying a constant theoretical maximum annual growth rate of 2%, indicates positive growth of the population in those areas where the electrocution threat from power lines is low, whereas the population in those areas where this threat is high is predicted to crash to extinction, from electrocution mortality alone, within a 20–35 year period. The regional population is predicted to show positive growth over the 50 year period. However, for a number of reasons that relate to the nature of certain parameters used in the model, the simulations must be considered to be conservative, at best. In addition, other unnatural mortality factors (notably inadvertent poisoning, drowning in high-walled farm reservoirs, harvesting for the traditional medicine trade, local food shortage), which are additive to power line-related mortality have not been taken into account. Management recommendations aimed at obtaining an improved estimate of the mean annual mortality rate from power lines, and at ameliorating the impact of electrocutions on the regional Cape Vulture population, are briefly mentioned. These address the former by documenting ways to improve the quantity and quality of the field data, and the latter by identifying areas where urgnt action needs to be taken to reduce or avoid the electrocution of vultures, by mitigating extant ‘unsafe’ power line infrastructure, and by ensuring that that only ‘safe’ infrastructure is used for new power lines.

Spatial and temporal changes in group dynamics and range use enable anti-predator responses in African buffalo

The reintroduction of large predators provides a framework to investigate responses by prey species to predators. Considerable research has been directed at the impact that reintroduced wolves (Canis lupus) have on cervids, and to a lesser degree, bovids, in northern temperate regions. Generally, these impacts alter feeding, activity, and ranging behavior, or combinations of these. However, there are few studies on the response of African bovids to reintroduced predators, and thus, there is limited data to compare responses by tropical and temperate ungulates to predator reintroductions. Using the reintroduction of lion (Panthera leo) into the Addo Elephant National Park (AENP) Main Camp Section, South Africa, we show that Cape buffalo (Syncerus caffer) responses differ from northern temperate ungulates. Following lion reintroduction, buffalo herds amalgamated into larger, more defendable units; this corresponded with an increase in the survival of juvenile buffalo. Current habitat preference of buffalo breeding herds is for open habitats, especially during the night and morning, when lion are active. The increase in group size and habitat preference countered initial high levels of predation on juvenile buffalo, resulting in a return in the proportion of juveniles in breeding herds to pre-lion levels. Our results show that buffalo responses to reintroduced large predators in southern Africa differ to those of northern temperate bovids or cervids in the face of wolf predation. We predict that the nature of the prey response to predator reintroduction is likely to reflect the trade-off between the predator selection and hunting strategy of predators against the life history and foraging strategies of each prey species.

An assessment of diet overlap of two mesocarnivores in the North West Province, South Africa

We used scat analysis to study the diet of two sympatric medium-sized carnivores: brown hyaena and black-backed jackal, in the North West Province of South Africa. Seven major dietary categories were identified from the scats, with mammal remains being most common for both species. Brown hyaena scats contained more large mammal remains, which together with the presence of invertebrates (in 50% of all brown hyaena scats), suggests that they mainly scavenged. Jackal scats contained a higher proportion of small mammal remains, suggesting that jackals actively hunted more often than brown hyaenas did. The diets differed significantly between the two species, even though diet overlap was fairly high (0.79). Further analysis, albeit based on small sample sizes, suggests that diet of these mesopredators differ between protected reserves with apex predators and unprotected areas without apex predators, thus confounding generalizations. Further studies are therefore required to investigate possible mesopredator release when apex predators are absent.