Anthrax outbreak among Grevy's zebra (Equus grevyi) in Samburu, Kenya

An anthrax outbreak occurred in the Wamba area of southern Samburu, Kenya, between December 2005 and March 2006. The outbreak affected equids including the endangered Grevy's zebras (Equus grevyi), plain zebras (Equis Burchelli) and donkeys (Equus asinus). Most of the deaths were localized in Nkaroni area just west of Wamba town. The diagnosis of anthrax was rapidly confirmed by bacteriological methods. The relevant government departments, including the Kenya Wildlife Service and Veterinary Department, and other stakeholders were promptly informed. Fifty‐three Grevy's zebra and 26 plains zebras died from anthrax. An equal number (eighteen) of adult male and female Grevy's zebras succumbed to the disease. The outbreak affected immature and mature individuals equally. The dead plain zebras included fifteen adult females, two adult males and nine immature individuals. The Veterinary Department responded by vaccinating livestock while Kenya Wildlife Service vaccinated 620 Grevy's zebras within southern Samburu. Examination of sites at which carcasses of animals which succumbed to the disease were burnt, revealed that unsupervised burning did not eliminate anthrax spores in 42% of the cases (n = 14). There is an urgent need to incorporate strategic wildlife disease monitoring in the struggle to save Grevy's zebras and other endangered species.     

Effects of Single‐ and Mixed‐Species Group Composition on the Flight Initiation Distances of Plains and Grevy's Zebras

Zebras, as prey species, attend to the behavior of nearby conspecifics and heterospecifics when making decisions to flee from predators. Plains zebras (Equus quagga) and Grevy's zebras (E. grevyi) frequently form mixed‐species groups in zones where their ranges overlap in Kenya. Although anecdotal observations suggest that Plains zebras are more flighty around humans than Grevy's zebras are, this has not been empirically confirmed, and relatively little is known about how they may influence each other's flight behavior. We addressed these questions by examining the flight initiation distances (FIDs) of Plains and Grevy's zebras in single‐species and mixed‐species groups from an approaching human. One target individual per group was approached steadily on foot, with start distance, alert distance, and FID recorded from this target. Using start distance and alert distance separately as covariates, 22 Plains zebras in single‐species groups exhibited a significantly longer mean FID than 15 Grevy's zebras in single‐species groups. The FIDs of 7 Plains zebras and 5 Grevy's zebras tested in mixed‐species groups were virtually equivalent and intermediate to those of Plains and Grevy's zebras in single‐species groups, suggesting a bidirectional moderating influence of heterospecifics on risk assessment. This effect was most pronounced for Plains zebras in mixed‐species groups that exhibited an FID that was significantly shorter than that of Plains zebras in single‐species groups. Our findings underscore the importance of recognizing that related equids may be differently impacted by anthropogenic stress.     

Selective Defecation and Selective Foraging: Antiparasite Behavior in Wild Ungulates?

Selective defecation and selective foraging are two potential antiparasite behaviors used by grazing ungulates to reduce infection by fecal–oral transmitted parasites. While there is some evidence that domestic species use these strategies, less is known about the occurrence and efficacy of these behaviors in wild ungulates. In this study, I examined whether wild antelope use selective defecation and selective foraging strategies to reduce exposure to gastrointestinal nematode parasites. By quantifying parasite levels in the environment in relation to the defecation patterns of three species, dik‐dik (Madoqua kirkii), Grant's gazelle (Gazella granti), and impala (Aepyceros melampus), I found that nematode larval concentrations in pasture were higher in the vicinity of clusters of feces (dung middens) compared to single fecal pellet groups or dung‐free areas. In addition, experimental feeding trials in free‐ranging dik‐dik showed that individuals selectively avoided feeding near concentrations of feces. Given that increased parasite contamination was found in the immediate vicinity of fecal clusters, fecal avoidance could help reduce host consumption of parasites and may therefore be an effective antiparasite behavior for certain species. On the other hand, while the concentration of parasite larvae in the vicinity of middens coupled with host avoidance of these areas during grazing could reduce host contact with parasites, results showing a positive correlation between the number of middens in a habitat and larval abundance at control sites suggest that dung middens might increase and not decrease overall host exposure to parasites. If this is the case, dung midden formation may not be a viable antiparasite strategy.     

Mammal road‐type associations in Kruger National Park,South Africa: Common mammals do not avoid tar roads more than dirt roads

The majority of Africa''s parks and conservation areas have a vast road network, facilitating motorized vehicle game viewing. These roads have an influence that is both road type‐ and species‐specific, on the surrounding ecosystem. Due to their higher traffic volumes, we hypothesized that tar roads and their immediate surrounds within the Kruger National Park, South Africa, are avoided to a greater extent by medium‐to‐large mammals than comparable dirt roads in the park. We systematically recorded the presence of medium‐to‐large mammal species from our vehicle, recording data at 401 tar and 369 dirt road stops in the Kruger National Park. In addition to species presence, we also estimated the proximity of animals to the road, as well as herd sizes. Our results indicate an equal likelihood of viewing the commonly recorded medium‐to‐large mammal species from both road types. The likelihood of observing larger herd sizes was also similar between tar and dirt roads for the three most commonly observed species, African elephant (Loxodonta africana), impala (Aepyceros melampus), and plains zebra (Equus quagga), and the likelihood of viewing impala and zebra close to the road also did not differ between tar and dirt roads. However, elephant was observed more often close to tar roads, compared to dirt roads. We interpreted this as the result of potentially increased woody cover associated with more water runoff in close proximity to tar roads compared with dirt roads. Our results not only have ecological significance, supporting the notion that many of the park''s species are habituated to human infrastructure, but also management implications, informing park officials about the influence of road traffic and road type on wildlife distributions.     

Fatal attraction: vegetation responses to nutrient inputs attract herbivores to infectious anthrax carcass sites

Parasites can shape the foraging behaviour of their hosts through cues indicating risk of infection. When cues for risk co-occur with desired traits such as forage quality, individuals face a trade-off between nutrient acquisition and parasite exposure. We evaluated how this trade-off may influence disease transmission in a 3-year experimental study of anthrax in a guild of mammalian herbivores in Etosha National Park, Namibia. At plains zebra (Equus quagga) carcass sites we assessed (i) carcass nutrient effects on soils and grasses, (ii) concentrations of Bacillus anthracis (BA) on grasses and in soils, and (iii) herbivore grazing behaviour, compared with control sites, using motion-sensing camera traps. We found that carcass-mediated nutrient pulses improved soil and vegetation, and that BA is found on grasses up to 2 years after death. Host foraging responses to carcass sites shifted from avoidance to attraction, and ultimately to no preference, with the strength and duration of these behavioural responses varying among herbivore species. Our results demonstrate that animal carcasses alter the environment and attract grazing hosts to parasite aggregations. This attraction may enhance transmission rates, suggesting that hosts are limited in their ability to trade off nutrient intake with parasite avoidance when relying on indirect cues.     

Bottom-up and top-down processes in African ungulate communities: resources and predation acting on the relative abundance of zebra and grazing bovids

African ungulate populations appear to be limited principally by their food resources. Within ungulate communities, plains zebras coexist with grazing bovids of similar body size, but rarely are the dominant species. Given the highly effective nutritional strategy of the equids and the resistance of zebras to drought, this is unexpected and suggests that zebra populations may commonly be limited by other mechanisms. Long-term research in the Serengeti ecosystem and in the Kruger National Park suggests that zebra could be less sensitive to food shortage, and more sensitive to predation, than grazing bovids: if this is a general principle, then, at a larger scale, resource availability should have a weaker effect on the abundance of zebra than on grazing ruminants of similar body size (wildebeest and buffalo), and zebras should be relatively more abundant in ecosystems where predators are rare or absent. We test these expectations using data on 23 near-natural ecosystems in east and southern Africa. The abundance of wildebeest is more closely related to resources than is that of zebra; buffalo are intermediate. We show that hyena densities are closely correlated with those of lions, and use the abundance of lions as an index of predation by large predators. The numerical response of lions to increases in the abundance of their prey was linear for mesoherbivores, and apparently so for the three species alone. Finally, the abundance of zebra relative to grazing bovids is lower in ecosystems with high biomasses of lions. These results indicate that zebras may commonly be more sensitive to top-down processes than grazing bovids: the mechanism(s) have not been demonstrated, but predation could play a role. If it is true, then when numbers of the large mammalian predators decline, zebra populations should increase faster than buffalo and wildebeest.     

The forgotten Grevy's zebra Equus grevyi population along the Kasigau Corridor ranches,SE Kenya: recent records and conservation issues

Since the 1970s, the globally endangered Grevy's zebra Equus grevyi has suffered a substantial reduction in range and population size. Grevy's zebra was introduced into the Tsavo ecosystem in two translocations: 22 individuals released in Tsavo East National Park in 1964 and 30 individuals released in Tsavo West National Park in 1977. This study focuses on the Tsavo East subpopulation within the Kasigau Corridor REDD+ Project Area. Data were collected from 2011 to 2014 using systematic and ad lib methods, while vegetation stratification was based on tonnes of CO₂e per hectare. Cumulatively, 785 Grevy's zebra were seen in 210 encounters. The largest single group comprised of 24 individuals; mean group size was 4 ± 3. Foals comprised 10% of all individuals seen and were recorded across most months, suggesting year‐round breeding. The range core for the Grevy's zebra in this area was estimated to be 400 km². Its density was estimated at 0.09 ± 0.038 individuals per km², translating to a population of 36 ± 15 individuals (range: 6–67) based on the estimated core range. Grasslands and sparse Acacia–Commiphora vegetation strata were used significantly more than expected. Despite constituting only about 2–3% of the global Grevy's zebra population, this subpopulation could still play a role as a – disconnected – reserve population.     

Effects of lions on behaviour and endocrine stress in plains zebras

Living under predation risk may alter both behaviour and physiology of potential prey. In extreme cases, such alterations may have serious demographic consequences, and recent studies support that non‐lethal effects of predation may have broad ecological consequences. However, behavioural and physiological responses to predation risk may be related to trade‐offs associated with resource acquisition and direct predation risk. We validated an enzyme‐linked immunoassay (EIA) for non‐invasive monitoring of stress in plains zebras (Equus quagga) from faecal material. We used this assay in combination with behavioural data to assess if plains zebras living with and without lions (Panthera leo) in a mountain savannah in southern Africa differed in behaviour and physiology, and if such differences were influenced by seasons with contrasting resource availability. Zebra group sizes did not differ between areas with and without lions, but zebra groups had more juveniles in an area with lions than groups in an area without lions, but only during the wet season. Similarly, we observed differences in individual vigilance, foraging behaviour and stress hormone concentrations, but all these differences were influenced by seasons. Despite these seasonal influences, our study did not suggest that zebras in an area with lions spent a higher proportion of time being vigilant, a lower proportion of time foraging, or had higher stress hormone levels. Our results instead suggest that zebras' responses to lion presence were highly context dependent and the result of complex interactions between resource abundance and cues about predation risk. Because of the obvious ecological and evolutionary ramifications of such findings, we argue that further research is needed to define the spatial and temporal scales over which predators impose indirect effects on their prey.     

Social structure,vigilance and behaviour of plains zebra (<Emphasis Type="Italic">Equus burchellii</Emphasis>): a 5-year case study of individuals living on a managed wildlife reserve

Most studies of plains zebra (Equus burchellii) have focused on population ecology and have not included long-term observations of identified individuals. Over a 5-year period, we studied the crepuscular activities of 13 individual zebras within a focal group held within a managed game reserve. We also examined individual residency within the group by recording births, mortalities and longevity of group membership by adults. Residency of individuals living in other groups on the reserve was similarly monitored to examine variability in social structure within this closed population over an extended period of time. Stable, female groups were the mainstay of group sociality with male mean residency at 31.6 months being variable in length or even absent. Social interactions across all categories of zebras were free from aggression. Despite an absence of non-human predators, the proportion of dusk time budget allocated to vigilance was high, at 41% for males during periods when they accompanied stable female groups and 12% for females during these same periods. Female vigilance increased significantly to 19% when males were not resident. Females spent 70% of the time grazing and males just 36%. Due to its long-term nature, we concluded this study established a base line for plains zebra activity that could assist in understanding the factors that influence the successful management and conservation of healthy populations.     

Antibodies against equine herpesviruses and equine arteritis virus in Burchell's zebras (Equus burchelli ) from the Serengeti ecosystem

A total of 51 sera from a migratory population of Burchell's zebras (Equus burchelli) were collected in the Serengeti National Park (Tanzania) between 1999 and 2001 to assess levels of exposure to equine herpesvirus types 1, 2, 4, 9 (EHV-1, -2, -4, -9), EHV-1 zebra isolate T965, and equine arteritis virus (EAV). Using virus-specific neutralizing antibody tests, seroprevalence was high for EHV-9 (60% of 45), moderate for EAV (24% of 51), and lower for the EHV-1-related zebra isolate (17% of 41), EHV-1 (14% of 49), and EHV-4 (2% of 50). No evidence for exposure to EHV-2 was found (0% of 51). The high level of exposure to EHV-9 is interesting because evidence of infection with this virus has not been previously described in any wild equine population. Although the epidemiology of EHV-9 in Burchell's zebras is presently unknown, our results suggest that in East Africa, this species may be a natural host of EHV-9, a neuropathogenic virus that was only recently isolated from captive Thomson's gazelles (Gazella thomsoni) in Japan. There is currently no evidence that EHV-9 induced mortality in Burchell's zebras in the Serengeti, but because of the reported virulence of this virus for more susceptible species such as Thomson's gazelles, viral transmission from infected zebras to ungulates may result in mortality.     

Habitat choice of Grevy's zebras (Equus grevyi) in Laikipia, Kenya

Characterizing habitat choice is essential for endangered species conservation. For the endangered Grevy's zebra ( Equus grevyi ), as with many widely ranging vertebrates, human activities may be an important factor affecting space use. Grevy's zebras are grazing ungulates inhabiting the savannahs of central-northern Kenya and Ethiopia. Past research on their social organization indicates that reproductive status shapes associations and movements. Here, we examine how habitat use varies across four reproductive classes: lactating and nonlactating females, bachelors and territorial males. We also test whether Grevy's zebra avoid locations close to active livestock corrals, or bomas . We find that forage quality, forage quantity and habitat openness of locations used by Grevy's zebra vary significantly depending on individual reproductive state. Lactating females and bachelors use areas with green, short grass and medium-dense bush more frequently than nonlactating females or territorial males. We hypothesize that lactating females trade off forage quantity and safety to access nutrients in growing grass. Across reproductive classes, Grevy's zebra choose locations further from active bomas than if they used the area randomly. Our results suggest that Grevy's zebra may require a range of vegetation characteristics for different reproductive classes. Further, they may need areas free from competition or disturbance by livestock.     

Nonhost species reduce parasite infection in a focal host species within experimental fish communities

The dilution effect describes the negative association between host biodiversity and the risk of infectious disease. Tests designed to understand the relative roles of host species richness, host species identity, and rates of exposure within experimental host communities would help resolve ongoing contention regarding the importance and generality of dilution effects. We exposed fathead minnows to infective larvae of the trematode, Ornithodiplostomum ptychocheilus in minnow‐only containers and in mixed containers that held 1–3 other species of fish. Parasite infection was estimated as the number of encysted worms (i.e., brainworms) present in minnows following exposure. The results of exposure trials showed that nonminnow fish species were incompatible with O. ptychocheilus larvae. There was no reduction in mean brainworm counts in minnows in mixed containers with brook sticklebacks or longnose dace. In contrast, brainworm counts in minnows declined by 51% and 27% in mesocosms and aquaria, respectively, when they co‐occurred with emerald shiners. Dilution within minnow + shiner containers may arise from shiner‐induced alterations in minnow or parasite behaviors that reduced encounter rates between minnows and parasite larvae. Alternatively, shiners may act as parasite sinks for parasite larvae. These results highlight the role of host species identity in the dilution effect. Our results also emphasize the complex and idiosyncratic effects of host community composition on rates of parasite infection within contemporary host communities that contain combinations of introduced and native species.     

Genetic markers and sample types for pedigree reconstruction in Grevy's zebra (Equus grevyi)

Effective captive population management relies on maintaining accurate pedigree records in order to direct breeding programmes and maximise conservation of genetic diversity. Where familial relationships or individual identities are unknown, it is now possible to recover this information via genetic analysis. We describe the first application of DNA profiling to reconstruct family groups of the Grevy's zebra (Equus grevyi), in order to identify three unknown stallions. To examine the potential for using non-invasive sampling techniques, DNA recovered from faeces, hair and blood was compared. Microsatellite loci derived from horses were tested for their utility in Grevy's zebra and a set of loci were subsequently employed to successfully identify the three animals via parental exclusion. The study demonstrates the relative ease with which DNA profiling can now be applied to captive management and highlights methods of genetic sampling that minimise stress to the animal and reduce sampling costs.     

Habitat use and movements of plains zebra (Equus burchelli) in response to predation danger from lions

Prey species must adapt their behavior to avoid predation. Asa key prey item for lions (Panthera leo), plains zebras (Equusburchelli) were expected to respond to immediate threats posedby lions in their area. In addition, zebras were predicted toexhibit behavior tuned to reduce the potential for encounterswith lions, by modifying their movement patterns in the timesof day and habitats of greatest lion danger. We studied a populationof approximately 600 plains zebra living in Ol Pejeta Conservancy,Kenya. We found that zebra abundance on or near a grasslandpatch was lower if lions had also been observed on that patchduring the same day. Predation danger was highest in grasslandhabitat during the night, when lions were more active. Zebrasightings and global positioning system radio collar data indicatedthat zebras also reduced their use of grassland at night, insteadusing more woodland habitat. Zebras moved faster and took sharperturns in grassland at night. It is hypothesized that these moreerratic movements assist zebras in avoiding detection or captureby lions.     

Horse haemoglobin phenotyping by agarose gel isoelectric focusing comparison of Thoroughbreds with other Equidae*

By using isoelectric focusing in thin agarose slab gels 1049 Thoroughbred, 82 Nooit-gedachter, 45 Percheron and 244 horses of other breeds were examined. The numbers of other Equidae tested were 107 donkeys, 50 mules, 4 common zebras (Equus burchelli boehmi) and 8 mountain zebras (Equus zebra hartmannae). Phenotypic data are presented for all tested animals and gene frequencies are calculated for the horses.     

Effects of Global Positioning System Collar Weight on Zebra Behavior and Location Error

ABSTRACT Global Positioning System (GPS) collars are increasingly being used to study fine-scale patterns of animal behavior. Previous studies on GPS collars have tried to determine the causes of location error without attempting to investigate whether the accuracy of fixes provides a correspondingly accurate measure of the animal's natural behavior. When comparing 2 types of GPS collar, we found a significant effect of collar weight and fit on the rate of travel of plains zebra (Equus burchelli antiquorum) females in the Makgadikgadi, Botswana. Although both types of collar were well within accepted norms of collar weight, the slightly heavier collars (0.6% of total body mass [TBM]) reduced rate of travel by >50% when foraging compared with the collar that was 0.4% of TBM. Collar effect was activity specific, particularly interfering with grazing behavior; the effect was less noticeable when zebras crossed larger interpatch distances. We highlight that small differences in collar weight or fit can affect specific behaviors, limiting the extrapolation of fine-scaled GPS data. This has important implications for wildlife biologists, who hitherto have assumed that collars within accepted weight limits have little or no effect on animal movement parameters.     

Grass greenness and grass height promote the resource partitioning among reintroduced Burchell's zebra and blue wildebeest in southern Mozambique

Differences in the selection of habitat and specific dietary items support resource partitioning and coexistence of sympatric African grazing herbivores, such as zebra and wildebeest. In Maputo Special Reserve (MSR), southern Mozambique, these two species were extirpated during the civil war (1977–1992); since 2010, they have been reintroduced into the Reserve. Identifying the resource selection by reintroduced species and how these species coexist, while utilising the same resources, is both of ecological interest and important for the management of wildlife communities and parks. This is a key application of our research. Therefore, the present study investigated resource partitioning between Burchell's zebra (Equus burchelli, Smuts 1832) and blue wildebeest (Connochaetes taurinus, Burchell 1823) in the MSR. We conducted the study from July 2016 to June 2017. The data were collected by direct observation, driving the vehicle along the reserve's roads that covered the vegetation communities where zebras and wildebeest are known to commonly occur. The composition of the diet and specific features of the grass grazed by the two species, including greenness, height, and the number of stems, were assessed. The widely available grass, Aristida barbicollis, contributed most to the diet of both herbivores. The dietary overlap between the two herbivores was higher during the dry season (95%) than wet season (86%). Resources partitioning appears to be determined, principally, by the height and greenness of the grass, with the zebra grazing taller grass, which may facilitate the access of the wildebeest to the greener, lower proportion of the forage. That results follow the expectation that, among native herbivores, overlap in resource use is not expected based on evolutionary segregation.     

Dung avoidance as a possible mechanism in competition between wild and domestic ungulates: two experiments with chamois <Emphasis Type="Italic">Rupicapra rupicapra</Emphasis>

Wild ungulates often adjust spatial behaviour where they coexist with livestock. In European mountains, chamois Rupicapra rupicapra or R. pyrenaica commonly avoid alpine pastures used by domestic sheep. Mechanisms leading to competitive losing out of the wild species are not well understood, but mostly, resource competition is inferred and sometimes demonstrated. We hypothesised that chamois need to minimise the risk of intestinal parasite uptake and therefore would avoid pastures contaminated with sheep dung. We tested this in two experiments by contaminating feeding patches of wild and captive chamois with (parasite-free) sheep faeces. Wild chamois did not avoid grazing sites or plots on which sheep dung had been placed at a (low) density representing the commonly encountered situation in the Swiss Alps. Captive chamois strongly reduced browsing time on small trees given for food when the surroundings of the trees were sprayed with a watery faecal solution. We concluded that the odour signalled a potential high risk of parasite infection to the captive chamois, whereas the density of sheep dung on the grazing site of wild chamois was not high enough to be perceived as a risk. The need to minimise endoparasite uptake from faeces may therefore play a role in driving spatial behaviour of wild ungulates and could result in competitive imbalance between wild and domestic ungulates.     

Rolling pits of Hartmann’s mountain zebra (Zebra equus hartmannae) increase vegetation diversity and landscape heterogeneity in the Pre‐Namib

Microsites created by soil‐disturbing animals are important landscape elements in arid environments. In the Pre‐Namib, dust‐bathing behavior of the near‐endemic Hartmann''s mountain zebra creates unique rolling pits that persist in the landscape. However, the ecohydrological characteristics and the effects of those microsites on the vegetation and on organisms of higher trophic levels are still unknown. In our study, we characterized the soil grain size composition and infiltration properties of rolling pits and reference sites and recorded vegetation and arthropod assemblages during the rainy season of five consecutive years with different amounts of seasonal rainfall. We further used the excess green vegetation index derived from drone imagery to demonstrate the different green up and wilting of pits and references after a rainfall event. In contrast to the surrounding grassland, rolling pits had finer soil with higher nutrient content, collected runoff, showed a higher infiltration, and kept soil moisture longer. Vegetation in the rolling pits was denser, dominated by annual forbs and remained green for longer periods. The denser vegetation resulted in a slightly higher activity density of herbivorous arthropods, which in turn increased the activity density of omnivorous and predatory arthropods. In times of drought, the rolling pits could act as safe sites and refuges for forbs and arthropods. With their rolling pits, Hartmann''s mountain zebras act as ecosystem engineers, contributing to the diversity of forb communities and heterogeneity of the landscape in the Pre‐Namib.     

Effect of sheep grazing on seed circulation on the Loess Plateau

In grazing ecosystems, mature seeds fall directly to the soil to form the soil seed bank (SSB), or are ingested by grazing livestock to become part of the dung seed bank (DSB; i.e., seed circulation). Both the SSB and DSB form the basis for the natural regeneration of vegetation. However, little is known about the relationships between the SSB, DSB, and aboveground vegetation (AGV) community under different stocking rates (SRs). This study investigated the relationships between the SSB, seeds in Tan sheep (Ovis aries) dung, and AGV at different SRs (0, 2.7, 5.3, and 8.7 sheep ha^–1) in a semiarid region of the Loess Plateau in China. We found that Tan sheep grazing increased the species richness heterogeneity of grassland vegetation, and negatively influenced the density of AGV. Under natural conditions, 17 species from soil‐borne seeds and 10 species from Tan sheep dung germinated. There was low species similarity between the soil and DSBs and AGV. Sheep SR and the seed banks (soil and dung) were negatively correlated with AGV. Seeds are cycled from herbage to livestock to soil during cold season grazing; the seasonal nature of this seed dispersal is an adaptation to harsh, semiarid environments. Increased seed bank diversity under sheep grazing facilitates grassland regeneration on the Loess Plateau, similarly to other semiarid regions globally.