Borrowing from Peter to pay Paul: managing threatened predators of endangered and declining prey species.

Conservation policy and practice can sometimes run counter to their mutual aims of ensuring species survival. In Kenya, where threatened predators such as lion deplete endangered prey such as Grevy's zebra, conservation practitioners seek to ensure species success through exclusive strategies of protection, population increase and preservation. We found strong selection for the endangered Grevy's zebra by both lion and hyena on two small fenced conservancies in Kenya. Despite abundant diversity of available prey, Grevy's zebra were selected disproportionately more than their availability, while other highly available species such as buffalo were avoided. Lions were therefore not alone in presenting a credible threat to Grevy's zebra survival. Conservation practitioners must consider interlinked characteristics of prey selection, resource availability and quality, the interplay between carnivore guild members and landscape scale population trends performance in wildlife management decisions.

A ghost fence-gap: surprising wildlife usage of an obsolete fence crossing.

Wildlife fencing has become more prevalent throughout Africa, although it has come with a price of increased habitat fragmentation and loss of habitat connectivity. In an effort to increase connectivity, managers of fenced conservancies can place strategic gaps along the fences to allow wildlife access to outside habitat, permitting exploration, dispersal and seasonal migration. Wildlife can become accustomed to certain movement pathways and can show fidelity to these routes over many years, even at the path level. Our study site has three dedicated wildlife crossings (fence-gaps) in its 142 km perimeter fence, and we continuously monitor these fence-gaps with camera-traps. We monitored one fence-gap before and after a 1.49 km fence section was completely removed and 6.8 km was reconfigured to leave only a two-strand electric fence meant to exclude elephant and giraffe, all other species being able to cross under the exclusionary fence. The removal and reconfiguration of the fence effectively rendered this fence-gap (which was left in place structurally) as a "ghost" fence-gap, as wildlife now had many options along the 8.29 km shared border to cross into the neighboring habitat. Although we documented some decline in the number of crossing events at the ghost-gap, surprisingly, 19 months after the total removal of the fence, we continued to document the usage of this crossing location by wildlife including by species that had not been previously detected at this location. We discuss potential drivers of this persistent and counterintuitive behavior as well as management implications.

Seroprevalence of Equine Herpesviruses 1 and 9 (EHV-1 and EHV-9) in Wild Grévy's Zebra ( Equus grevyi) in Kenya.

Equid herpesviruses types 1 (EHV-1) and 9 (EHV-9) are unusual among herpesviruses in that they lack strong host specificity, and the full extent of their host range remains unclear. The virus establishes latency for long periods and can be reactivated and shed, resulting in clinical disease in susceptible species. A sensitive and specific peptide-based enzyme-linked immunosorbent assay was developed to study the seroprevalence of both viruses in a broad range of species among both wild and captive populations. We used this assay to study the seroprevalences of EHV-1 and EHV-9 in a natural population of the highly endangered Grévy's zebra ( Equus grevyi) in Kenya, sampled during a 4-yr period (2012-15). The results were compared with those obtained from captive Grévy's zebras from a previous study. The wild population had a significantly higher seroprevalence of EHV-9 compared with the captive population, suggesting that captivity might reduce exposure to this serotype. In contrast, the seroprevalences of EHV-1 between captive and wild groups was not significantly different. The seroprevalence of EHV-9 was not significantly higher than EHV-1 in zebras within the wild Kenyan population.

Testing the effects of perimeter fencing and elephant exclosures on lion predation patterns in a Kenyan wildlife conservancy.

The use of fences to segregate wildlife can change predator and prey behaviour. Predators can learn to incorporate fencing into their hunting strategies and prey can learn to avoid foraging near fences. A twelve-strand electric predator-proof fence surrounds our study site. There are also porous one-strand electric fences used to create exclosures where elephant (and giraffe) cannot enter in order to protect blocs of browse vegetation for two critically endangered species, the black rhinoceros (Diceros bicornis) and the Grevy's zebra (Equus grevyi). The denser vegetation in these exclosures attracts both browsing prey and ambush predators. In this study we examined if lion predation patterns differed near the perimeter fencing and inside the elephant exclosures by mapping the location of kills. We used a spatial analysis to compare the predation patterns near the perimeter fencing and inside the exclosures to predation in the rest of the conservancy. Predation was not over-represented near the perimeter fence but the pattern of predation near the fence suggests that fences may be a contributing factor to predation success. Overall, we found that predation was over-represented inside and within 50 m of the exclosures. However, by examining individual exclosures in greater detail using a hot spot analysis, we found that only a few exclosures contained lion predation hot spots. Although some exclosures provide good hunting grounds for lions, we concluded that exclosures did not necessarily create prey-traps per se and that managers could continue to use this type of exclusionary fencing to protect stands of dense vegetation.

Testing the Prey-Trap Hypothesis at Two Wildlife Conservancies in Kenya.

Protecting an endangered and highly poached species can conflict with providing an open and ecologically connected landscape for coexisting species. In Kenya, about half of the black rhino (Diceros bicornis) live in electrically fenced private conservancies. Purpose-built fence-gaps permit some landscape connectivity for elephant while restricting rhino from escaping. We monitored the usage patterns at these gaps by motion-triggered cameras and found high traffic volumes and predictable patterns of prey movement. The prey-trap hypothesis (PTH) proposes that predators exploit this predictable prey movement. We tested the PTH at two semi-porous reserves using two different methods: a spatial analysis and a temporal analysis. Using spatial analysis, we mapped the location of predation events with GPS and looked for concentration of kill sites near the gaps as well as conducting clustering and hot spot analysis to determine areas of statistically significant predation clustering. Using temporal analysis, we examined the time lapse between the passage of prey and predator and searched for evidence of active prey seeking and/or predator avoidance. We found no support for the PTH and conclude that the design of the fence-gaps is well suited to promoting connectivity in these types of conservancies.

Prevalence of Theileria equi and Babesia caballi as well as the identification of associated ticks in sympatric Grevy's zebras (Equus grevyi) and donkeys (Equus africanus asinus) in northern Kenya.

The role of equine piroplasmosis as a factor in the population decline of the Grevy's zebra is not known. We determined the prevalence of Babesia caballi and Theileria equi in cograzing Grevy's zebras (Equus grevyi) and donkeys (Equus africanus asinus) in northern Kenya and identified the associated tick vectors. Blood samples were taken from 71 donkeys and 16 Grevy's zebras from March to May 2011. A nested PCR reaction using 18s ribosomal (r)RNA primers on 87 blood spots showed 72% (51/71; 95% confidence interval [CI] 60.4-81.0%) of donkeys and 100% (16/16; 95% CI, 77.3-100%) of Grevy's zebras were T. equi positive. No samples were positive for B. caballi. Sequence comparison using the National Center for Biotechnology Information's basic local alignment search tool identified homologous 18s rRNA sequences with a global geographic spread. The T. equi-derived sequences were evaluated using Bayesian approaches with independent Metropolis-coupled Markov chain Monte Carlo runs. The sequences clustered with those found in Sudan, Croatia, Mongolia, and the US, with statistical support greater than 80% for the two main clades. Hyalomma tick species were found on both donkeys and Grevy's zebras, whereas Rhipicephalus pulchellus was found exclusively on Grevy's zebras and Hyalomma marginatum rupfipes on donkeys. The prevalence of T. equi was 100% in Grevy's zebras and 72% in donkeys with common tick vectors identified. Our results suggest that donkeys and Grevy's zebras can be asymptomatic carriers and that piroplasmosis is endemic in the study area.

Seasonal diet and prey preference of the African lion in a waterhole-driven semi-arid savanna.

Large carnivores inhabiting ecosystems with heterogeneously distributed environmental resources with strong seasonal variations frequently employ opportunistic foraging strategies, often typified by seasonal switches in diet. In semi-arid ecosystems, herbivore distribution is generally more homogeneous in the wet season, when surface water is abundant, than in the dry season when only permanent sources remain. Here, we investigate the seasonal contribution of the different herbivore species, prey preference and distribution of kills (i.e. feeding locations) of African lions in Hwange National Park, Zimbabwe, a semi-arid African savanna structured by artificial waterholes. We used data from 245 kills and 74 faecal samples. Buffalo consistently emerged as the most frequently utilised prey in all seasons by both male (56%) and female (33%) lions, contributing the most to lion dietary biomass. Jacobs' index also revealed that buffalo was the most intensively selected species throughout the year. For female lions, kudu and to a lesser extent the group "medium Bovidae" are the most important secondary prey. This study revealed seasonal patterns in secondary prey consumption by female lions partly based on prey ecology with browsers, such as giraffe and kudu, mainly consumed in the early dry season, and grazers, such as zebra and suids, contributing more to female diet in the late dry season. Further, it revealed the opportunistic hunting behaviour of lions for prey as diverse as elephants and mice, with elephants taken mostly as juveniles at the end of the dry season during droughts. Jacobs' index finally revealed a very strong preference for kills within 2 km from a waterhole for all prey species, except small antelopes, in all seasons. This suggested that surface-water resources form passive traps and contribute to the structuring of lion foraging behaviour.

Understanding patch departure rules for large carnivores: lion movements support a patch-disturbance hypothesis.

We test two hypotheses that could account for patch departure by large mammalian carnivores. One hypothesis is the unsuccessful-hunt hypothesis, where carnivores leave an area after an unsuccessful hunt but continue hunting in the same area after a successful hunt. The second hypothesis is the patch-disturbance hypothesis, where carnivores depart the area after a successful hunt because of behavioral responses of prey to predator presence. We used global positioning system collars to monitor the movements of African lions (Panthera leo) and identified their kill sites to distinguish between these two hypotheses. Lions moved to a different area (≥ 5 km away) after 87% of the kills, which supports the patch-disturbance hypothesis for patch-departure behavior of large mammalian carnivores.