Divergent water requirements partition exposure risk to parasites in wild equids

For grazing herbivores, dung density in feeding areas is an important determinant of exposure risk to fecal‐orally transmitted parasites. When host species share the same parasite species, a nonrandom distribution of their cumulative dung density and/or nonrandom ranging and feeding behavior may skew exposure risk and the relative selection pressure parasites impose on each host. The arid‐adapted Grevy''s zebra (Equus grevyi) can range more widely than the water‐dependent plains zebra (Equus quagga), with which it shares the same species of gastrointestinal nematodes. We studied how the spatial distribution of zebra dung relates to ranging and feeding behavior to assess parasite exposure risk in Grevy''s and plains zebras at a site inhabited by both zebra species. We found that zebra dung density declined with distance from water, Grevy''s zebra home ranges (excluding those of territorial males) were farther from water than those of plains zebras, and plains zebra grazing areas had higher dung density than random points while Grevy''s zebra grazing areas did not, suggesting a greater exposure risk in plains zebras associated with their water dependence. Fecal egg counts increased with home range proximity to water for both species, but the response was stronger in plains zebras, indicating that this host species may be particularly vulnerable to the elevated exposure risk close to water. We further ran experiments on microclimatic effects on dung infectivity and showed that fewer nematode eggs embryonated in dung in the sun than in the shade. However, only 5% of the zebra dung on the landscape was in shade, indicating that the microclimatic effects of shade on the density of infective larvae is not a major influence on exposure risk dynamics. Ranging constraints based on water requirements appear to be key mediators of nematode parasite exposure in free‐ranging equids.     

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.     

Treatment of equine sarcoid in seven Cape mountain zebra (Equus zebra zebra)

Equine sarcoid has been diagnosed in endangered Cape mountain zebra (Equus zebra zebra) in at least two game reserves in South Africa, with prevalence as high as 53% in Bontebok National Park. Seven Cape mountain zebras with sarcoids were treated with either surgical excision, 5-fluorouracil, allogenous vaccine, or a combination of 5-fluorouracil and allogenous vaccine. One of the two sarcoids on one of the 5-fluorouracil-treated zebras was left untreated. The microscopic features of the tumors evaluated showed either all or most of the typical epidermal and dermal histologic features of equine sarcoid. The zebras were examined 2 yr posttreatment to determine outcome. All sarcoids had resolved except on the zebra on which one of the sarcoids was left untreated. The efficacy of the three treatment methods in Cape mountain zebra is encouraging.     

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.     

Further evidence for sex differences in suckling behaviour of captive plains zebra foals

Sex differences in suckling behaviour could be explained either by different maternal investment or by different behaviour of young. Previously, we found sex differences in rejection and termination of suckling bouts in captive plains zebras (Equus burchellii). In this study, we analysed the length of suckling bout interruptions and unsuccessful subsequent attempts which followed a successful suckling bout in plains zebra foals at Dvůr Králové Zoo. When a suckling bout was interrupted, male foals resumed the bout sooner than female foals. The probability of a subsequent attempt decreased with increasing age in female foals but did not change in male foals. These results demonstrate higher effort in suckling interest by male foals, who tried to suckle more than female foals. Although we found sex bias in the effort to suckle, which suggested higher parent–offspring conflict in male offspring, we cannot omit a sex bias in milk supply by the mother.     

Tear production in three captive wild herbivores in Israel

The Schirmer tear test (STT) I was performed to evaluate tear production in 12 captive Nubian ibex (Capra ibex nubiana), 10 captive Burchell's zebras (Equus burchelli) and five Arabian oryx (Oryx leucoryx) at the Tel-Aviv Ramat-Gan Zoological Center (Israel). Mean (+/- standard deviation) STT values were 13.2 +/- 5.1 mm/min in the ibex, 23.4 +/- 3.4 mm/min in the zebra and 12.7 +/- 4.8 mm/min in the oryx. There were no significant effects of gender, age, weight, or side of the eye. There were no significant differences in STT values between ibex and oryx, but tear production in both species was significantly lower than in zebras. Knowledge of normal tear production values is important for the differential diagnosis of conjunctivitis and keratitis in these species.     

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.     

Clinico-Pathologic Findings in Young Pigs Fed Different Levels of Selenium,Vitamin E and Antioxidant

A randomized, blocked 23 factorial experiment was conducted with 48 young pigs. The treatment factors were: 2 levels of selenium (55 and 115 µg/kg), 2 levels of vitamin E (3 and 53 mg/kg) and 2 levels of the antioxidant feed additive Ethoxyquin (0 and 150 mg/kg). All pigs were kept in single pens and fed ad libitum throughout the experimental period of 9 weeks, i.e. from 3 to 12 weeks of age. Plasma, heart, liver and muscle Se levels as well as whole blood glutathione peroxidase activity (EC 1.11.1.9 GSH-Px) were significantly higher in pigs given a dietary supplement of Se than in pigs given no supplement of Se (P ≤ 0.001). The Se-supplemented pigs showed a tendency to lower mean serum transaminase activity (ASAT and ALAT) than unsupplemented pigs, but the influence was significant (P ≤ 0.05) only for the ALAT activity. Blood vit. E levels were higher for pigs receiving a supplement of vit. E than for unsupplemented pigs (P ≤ 0.001), and so was the resistance of red blood cells against lipid peroxidation (ELP), as expressed by lower ELP values. There were no effects of Ethoxyquin supplementation on the biochemical variables included in the study. The histological examination of heart muscle showed that the score for changes was negatively influenced by both Se and vit. E supplement (P ≤ 0.001) and to some extent also by Ethoxyquin supplement (P ≤ 0.05). The histological picture of m. long dorsi was influenced only by the vit. E supplement (P ≤ 0.01). No histological changes were found in the liver in this study. There were inverse relationships between whole blood GSH-Px defluorescence time and blood Se, and between ELP and whole blood vit. E (P ≤ 0.001).     

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.     

A rapid loss of stripes: the evolutionary history of the extinct quagga

Twenty years ago, the field of ancient DNA was launched with the publication of two short mitochondrial (mt) DNA sequences from a single quagga (Equus quagga) museum skin, an extinct South African equid (Higuchi et al. 1984 Nature312, 282-284). This was the first extinct species from which genetic information was retrieved. The DNA sequences of the quagga showed that it was more closely related to zebras than to horses. However, quagga evolutionary history is far from clear. We have isolated DNA from eight quaggas and a plains zebra (subspecies or phenotype Equus burchelli burchelli). We show that the quagga displayed little genetic diversity and very recently diverged from the plains zebra, probably during the penultimate glacial maximum. This emphasizes the importance of Pleistocene climate changes for phylogeographic patterns in African as well as Holarctic fauna.     

Comparative Effect of Selenium in Wheat,Barley, Fish Meal and Sodium Selenite for Prevention of Exudative Diathesis in Chicks

Groups af White Leghorn chicks obtained from dams deprived on selenium (Se), were fed from hatching a low-Se-vitamin E basal diet alone, or supplemented with 0.02, 0.04, 0.06 or 0.08 mg Se/kg diet, as sodium selenite (Na2SeO3 · 5H2O), wheat, barley or fish meal. Prevention of the Se-vitamin E deficiency responsive disease exudative diathesis (ED) as it was clinical observed, induction of the plasma Se dependent enzyme glutathione peroxidase (GSH-Px) activity, and Se concentration in the cardiac muscle were observed to be dietary Se level and source dependent. Slope ratio assay was applied to estimate the biological availability of Se in the natural sources relative to Se in sodium selenite. For the prevention of ED, the bioavailability of Se in wheat, barley and fish meal was 99, 85 and 80 %, respectively. The increase in the plasma GSH-Px activity revealed a bioavailability for Se in wheat, barley and fish meal of 79, 71 and 66 %, respectively. Using retention of Se in the cardiac muscle as the bioassay, a bioavailability of 108, 87 and 100 % was calculated for wheat, barley and fish meal Se, respectively.     

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.     

Evidence for altered structure and impaired mitochondrial electron transport function in selenium deficiency

Selenium (Se) deficiency in the experimental models,Coturnix coturnix japonica andCorcyra cephalonica, resulted in impaired mitochondrial substrate oxidations and lowered thiol levels. Studies with respiratory inhibitors confirmed reduced mitochondrial electron transport enzyme activities, especially at cytochromec oxidase (COX), the terminal segment. Enhanced mitochondrial lipid peroxidation in Se deficiency was more pronounced in the heart tissue of the quail compared to other tissues. Glutathione peroxidase (GSH-Px) activity toward H₂O₂ and cumene hydroperoxide were generally low in the insect muscle tissue and activity toward H₂O₂ was maximal in the quail heart mitochondria that was not very sensitive to Se status. Lowered COX activity in Se deficiency was more directly correlated with the increased level of lipid peroxidation than with the GSH-Px activity measured, suggestive of Se mediated protective mechanisms independent of GSH-Px. Electron microscopic observations revealed structural changes such as loss of cristae with proliferative and degenerative changes of the mitochondria in Se deficiency. Involvement of Se in maintaining structure and functional efficiency of mitochondria is evident from the present study.     

Reactive responses of zebras to lion encounters shape their predator–prey space game at large scale

The predator–prey space game and the costs associated with risk effects are affected by prey 1) proactive adjustments (when prey modify their behaviour in response to an a priori assessment of the risk level) and 2) reactive adjustments (when prey have detected an immediate threat). Proactive adjustments are generally well‐studied, whereas the frequency, strength and duration of reactive adjustments remain largely unknown. We studied the space use and habitat selection of GPS‐collared zebras Equus quagga from 2 to 48 h after an encounter with lions Panthera leo. Lion–zebra encounters generally occurred close to artificial waterholes (< 1 km). Two hours after an encounter, zebras were more likely to have fled than stay when the encounter occurred in more risky bushy areas. During their flight, zebras selected grasslands more than usual, getting great visibility. Regardless of their initial response, zebras finally fled at the end of the night and reached areas located far from waterholes where encounters with lions are less frequent. The large‐scale flights (～4–5 km) of zebras led to a local zebra depression for lions. Zebras that had fled immediately after the encounter resumed their behaviour of coming close to waterholes on the following day. However, zebras that had initially stayed remained far from waterholes for an extra 24 h, remaining an elusive prey for longer. The delay in the flight decision had different short‐term consequences on the lion–zebra game. We reveal that the spatial context of the encounter shapes the immediate response of prey, and that encountering predators induces strong behavioural responses: prey flee towards distant, safer, areas and have a constrained use of key resource areas which are at the heart of the predator–prey game at larger spatio‐temporal scales. Nighttime encounters were infrequent (once every 35 days on average), zebra responses were short‐lived (< 36 h) but occurred over a large spatial scale (several km).     

Effects of Dietary Selenium Deficiency or Excess on Gene Expression of Selenoprotein N in Chicken Muscle Tissues

Previous studies have determined the effects of dietary selenium (Se) supplementation on selenoprotein N (SelN, SEPN1), selenophosphate synthetase-1 (SPS1), and selenocysteine-synthase (SecS) mRNA abundance in chicken skeletal and cardiac muscles. To investigate collective responses of these genes to dietary Se concentrations ranging from deficiency to moderately high level in muscle tissues of chicken, 1-day-old chickens were exposed to a diet of deficient Se and supplemented with Se (0.15 mg Se/kg and 1.50 mg Se/kg) as sodium selenite in the feed for 35 days. Muscle tissues (flight, breast, leg, and cardiac muscles) were collected and examined for Se content and mRNA levels of SelN on days 1, 15, 25, and 35 days, respectively. Moreover, SPS1 and SecS mRNA levels were analyzed. The results showed that the expression of SelN gene in cardiac muscle responded to dietary Se concentrations. SelN gene was downregulated in the Se deficiency group (L group), and upregulated in the Se excess group (H group) compared with the moderate Se group (M group) (P?<?0.05) in cardiac muscle. Se deficiency mainly unregulated SelN mRNA level in skeletal muscles compared with M group. Excess dietary Se mainly resulted in the upregulation of SelN mRNA level in skeletal muscles compared with the M group. SecS mRNA levels responded to dietary Se concentrations showed a similar change compared with SelN in cardiac muscle. SPS1 mRNA levels responded to dietary Se concentrations showed a downregulation in L group and upregulation in H group. However, SelN mRNA levels displayed a different expression pattern in different skeletal and cardiac muscles. Moreover, Se also regulated the levels of SPS1 and SecS mRNAs. In summary, Se regulated the expression of SelN gene and affected the mRNA levels of SecS and SPS1. The level of Se in the feed may regulate SelN biosynthesis by affecting the levels of SPS1 and SecS mRNA.     

Significance of alterations in hepatic antioxidant enzymes. Primacy of glutathione peroxidase.

The relative contributions of catalase and the selenoenzyme glutathione peroxidase (GSH-Px) were elucidated in the rat liver by selectively modulating the activities of these enzymes using dietary selenium (Se) and the catalase inhibitor 3-amino-1,2,4-triazole (3-AT). Increased peroxidation occurred only in Se-deficient rats with markedly reduced cytosolic and mitochondrial GSH-Px activities. Although 3-AT treatment resulted in a 75% reduction of hepatic catalase activity and also a 20% reduction of both cytosolic and mitochondrial superoxide dismutase (SOD) activity, no incremental increase in peroxidation was observed over that associated with Se deficiency. In Se-deficient animals, treatment with 3-AT resulted in a doubling of cytosolic GSH-Px. This was associated with a 49% elevation in hepatic Se suggesting that increased Se may have contributed to the enhanced GSH-Px activity. These results suggest that GSH-Px plays the pivotal role in preventing hepatic peroxidation. Furthermore, the effects of 3-AT in vivo are not restricted to inhibition of catalase activity insofar as it also affects cytosolic GSH-Px activity and cytosolic and mitochondrial SOD activities.     

Influence of selenium deficiency on vital functions in rats

To clarify the relationship between selenium (Se) deficiency and functional disorders, the authors determined the Se concentration, anti-oxidant enzyme activity, and other parameters in rats fed a Se-deficient diet. Rats fed the Se-deficient diet showed a decrease in Se concentration and glutathione peroxidase (GSH-Px) activity in plasma, erythrocytes, heart, liver, and skeletal muscle from the first week after the initiation of the diet, an increase in heart lipid peroxide concentration from the second week, and an increase in liver glutathione S-transferase activity from the fourth week. From the twelfth week, a decrease in the growth rate in the rats fed the Se-deficient diet was observed. In spite of this growth impairment, no changes in electrocardiogram, muscle tone, degree of hemolysis, plasma biochemistry, or hematological values were detected. In summary, the authors found that a reduction of body Se is easily induced, but that the appearance of functional disorders following Se deficiency is difficult to detect in rats.     

What limits the Serengeti zebra population?

The populations of the ecologically dominant ungulates in the Serengeti ecosystem (zebra, wildebeest and buffalo) have shown markedly different trends since the 1960s: the two ruminants both irrupted after the elimination of rinderpest in 1960, while the zebras have remained stable. The ruminants are resource limited (though parts of the buffalo population have been limited by poaching since the 1980s). The zebras resource acquisition tactics should allow them to outcompete the ruminants, but their greater spatial dispersion makes them more available to predators, and it has been suggested that this population is limited by predation. To investigate the mechanisms involved in the population dynamics of Serengeti zebra, we compared population dynamics among the three species using demographic models based on age-class-specific survival and fecundity. The only major difference between zebra and the two ruminants occurred in the first-year survival. We show that wildebeest have a higher reproductive potential than zebra (younger age at first breeding and shorter generation time). Nevertheless, these differences in reproduction cannot account for the observed differences in the population trends between the zebra and the ruminants. On the other hand, among-species differences in first-year survival are great enough to account for the constancy of zebra population size. We conclude that the very low first-year survival of zebra limits this population. We provide new data on predation in the Serengeti and show that, as in other ecosystems, predation rates on zebras are high, so predation could hold the population in a predator pit. However, lion and hyena feed principally on adult zebras, and further work is required to discover the process involved in the high mortality of foals.     

Comparison of selenium level and glutathione peroxidase activity in tissues of vitamin B6-deficient rats fed sodium selenite or DL-selenomethionine

The effect of vitamin B₆ on the levels of tissue selenium (Se) and glutathione peroxidase (GSH-Px) was studied. Male Wistar 4-week-old rats were fed a vitamin B₆-Se-deficient basal diet for 2 weeks, then divided into 10 groups of five or six rats and fed their respective diets for 4 weeks. The experimental design was a 2×2×2 factorial with two levels of vitamin B₆, two forms of Se, and two levels of Se, plus two extra groups (vitamin B₆-supplemented and deficient without Se). Vitamin B₆ was 0 and 250 μg pyridoxine-HCl/100 g of diet; Se forms were Na₂SeO₃ and DL-selenomethionine; Se levels were 0.5 and 5.0 mg Se/kg of diet. Regardless of form or level of Se, vitamin B₆-deficient rats had lower body weights and organ weights than vitamin B₆-supplemented rats. At 5.0 mg Se/kg of diet, Na₂SeO₃ caused a further depression. Vitamin B₆ deficiency resulted in a higher Se level and GSH-Px activity in plasma of rats fed selenomethionine. However, Se content an GSH-Px activity in erythrocytes were significantly elevated in vitamin B₆-supplemented rats compared with vitamin B₆-deficient rats. Se levels in muscle and heart were significantly lower in vitamin B₆-deficient groups fed Na₂SeO₃ than in vitamin B₆-supplemented groups. Vitamin B₆-deficient rats fed selenomethionine had higher Se levels in muscle, heart, spleen, liver, and kidneys than vitamin B₆-supplemented rats. Activity of GSH-Px in muscle, heart, and spleen was significantly lower in vitamin B₆-deficient groups than in vitamin B₆-supplemented groups, regardless of form of Se. A significant decrease of GSH-Px in liver was observed in vitamin B₆-deficient rats fed selenomethionine compared with vitamin B₆-supplemented rats, whereas no significant decrease was observed in those fed Na₂SeO₃. These results suggest that vitamin B₆ is involved in the distribution and transportation of Se in body and the metabolism of selenomethionine in liver.     

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.