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).     

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.     

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.     

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.     

Clinical and Pathologic Features of a Suspected Selenium Deficiency in Captive Plains Zebras

Previous studies have shown that selenium (Se) deficiency is associated with nutritional myopathy, known as white muscle disease (WMD), in horses. However, correlations between Se deficiency and clinical findings, such as hematologic biochemical values and pathological features, have not been evaluated in captive plains zebras. The purpose of the present study was to investigate the clinical and pathologic features that may be caused by a Se deficiency in the captive plains zebra. Clinical findings, feed analyses, hematologic biochemical analyses, response to treatment, and pathologic examination were assessed in six affected plains zebras. The dietary concentration of Se in feed was also tested. Sudden death occurred in two cases during the first day of the onset of symptoms. Two zebras died at 4 days and two zebras survived after treatment. The clinical signs in affected animals were characterized by general weakness, astasia, and abnormal postural positions. The Se concentration in hay from the breeding stable was low, based on the reference value. Glutathione peroxidase (GSH-Px) activity was lower compared with the equine reference value. Multiple areas of subcutaneous steatitis and pale skeletal muscle and myocardium were revealed at gross necropsy. Degeneration and necrosis of myocardial and skeletal muscles, as well as congestion of the liver, lung, and kidney were found via histopathological examination. No suspected bacterial infections were found. Feed analyses, response to treatment, serum GSH-Px activity, and pathological features suggest that Se deficiency may have caused the disease in the six affected captive plains zebra.     

Profitability, encounter rates, and prey choice of African lions

The prey preferences of African lions (Panthera leo) in SerengetiNational Park, Tanzania, were examined in three ways. First,lion encounter rates with prey types were measured and comparedwith a random sample of the prey population. Lions encounteredmore wart hogs (Phacochoerus aethiopicus), Grant's gazelles(Gazella granti), wildebeests (ConnochaeUs taurinus), and zebras(Equus burchelli) than expected. Second, preferred prey typesof lions were identified using conditional logit analysis. Lionspreferred to hunt small prey groups, groups that were closerthan 200 m, and groups that contained wart hogs, wildebeests,or zebras. Third, a risk-minimization optimal foraging modeland a rate-maximization model were used to predict lion preferences.The foraging theory models predict that preferences should changewith season and with lion group size. Qualitative support wasfound for most of these predictions.     

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.     

Zebra Stripes through the Eyes of Their Predators,Zebras, and Humans

The century-old idea that stripes make zebras cryptic to large carnivores has never been examined systematically. We evaluated this hypothesis by passing digital images of zebras through species-specific spatial and colour filters to simulate their appearance for the visual systems of zebras’ primary predators and zebras themselves. We also measured stripe widths and luminance contrast to estimate the maximum distances from which lions, spotted hyaenas, and zebras can resolve stripes. We found that beyond ca. 50 m (daylight) and 30 m (twilight) zebra stripes are difficult for the estimated visual systems of large carnivores to resolve, but not humans. On moonless nights, stripes are difficult for all species to resolve beyond ca. 9 m. In open treeless habitats where zebras spend most time, zebras are as clearly identified by the lion visual system as are similar-sized ungulates, suggesting that stripes cannot confer crypsis by disrupting the zebra’s outline. Stripes confer a minor advantage over solid pelage in masking body shape in woodlands, but the effect is stronger for humans than for predators. Zebras appear to be less able than humans to resolve stripes although they are better than their chief predators. In conclusion, compared to the uniform pelage of other sympatric herbivores it appears highly unlikely that stripes are a form of anti-predator camouflage.     

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.     

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.     

Diet quality in a wild grazer declines under the threat of an ambush predator

Predators influence prey populations not only through predation itself, but also indirectly through prompting changes in prey behaviour. The behavioural adjustments of prey to predation risk may carry nutritional costs, but this has seldom been studied in the wild in large mammals. Here, we studied the effects of an ambush predator, the African lion (Panthera leo), on the diet quality of plains zebras (Equus quagga) in Hwange National Park, Zimbabwe. We combined information on movements of both prey and predators, using GPS data, and measurements of faecal crude protein, an index of diet quality in the prey. Zebras which had been in close proximity to lions had a lower quality diet, showing that adjustments in behaviour when lions are within short distance carry nutritional costs. The ultimate fitness cost will depend on the frequency of predator–prey encounters and on whether bottom-up or top-down forces are more important in the prey population. Our finding is the first attempt to our knowledge to assess nutritionally mediated risk effects in a large mammalian prey species under the threat of an ambush predator, and brings support to the hypothesis that the behavioural effects of predation induce important risk effects on prey populations.     

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.     

Evidence that the parasitic nematode Skrjabinoclava manipulates host Corophium behavior to increase transmission to the sandpiper, Calidris pusilla

We found evidence that a nematode (Skrjabinoclava morrisoni) adaptivelymanipulates the behavior of its intermediate host (the amphipod Corophiumvolutator) to increase its likelihood of transmission to itsfinal host (the semipalmated sandpiper, Calidris pusilla). We foundthat male and female amphipods parasitized by nematodes increasedtheir surface activity in the field during daytime, but notduring nighttime hours. Increased surface activity is knownto increase susceptibility of amphipods to predation by sandpipersduring the day, but not at night, when sandpipers do not feedvisually. Also, as predicted by the manipulation hypothesis,only late-stage (infective) larvae of nematodes were associatedwith behavioral changes of amphipods. We found no evidence thatparasites were associated with other amphipod behaviors in thelaboratory, such as trail complexity, distance traveled, orburrow-probing activity of crawling males as would be expectedif parasitized hosts altered their own behavior. Survivorshipof amphipods was also unaffected by parasitism, which may favorparasite transmission. Thus, behavioral changes of parasitizedhosts were simple, and their expression was context-dependentand related to likelihood of predation. We argue that maturationtimes of nematodes in relation to migration schedules of sandpipers providea narrow window of opportunity and may explain why nematodes manipulateamphipod behavior.     

Non-offspring nursing in social carnivores: minimizing the costs

We compare the nursing behavior of two species, African lions(PanfheraLeo) and spotted hyenas(Crocuta Craig Packer crocuta), and showthat non-offspring nursing is much less common in hyenas thanlions. Hyenas spend less time with their cubs, are more alertduring the suckling attempts of cubs, and more frequently resistthe attempts of non-offspring. Vigilance against milk theftmay therefore influence the distribution of non-offspring nursingacross species. Our detailed study of non-offspring nursingin lions shows that females preferentially nurse their own offspringand that cubs are more surreptitious when attempting to sucklefrom other females. Non-offspring nursing in lions is most commonwhen the costs are lowest. First, non-offspring nursing is morecommon among close kin. Second, females with small litters,and presumably more milk to spare, give a higher proportionof their nursing to non-offspring. Third, females give a higherproportion of their nursing to non-offspring as their own cubsgrow older and need less milk. Cubs reared in créchesdo not appear to gain more milk that cubs raised alone, andfemales do not show any evidence of reciprocity in nursing oneanother's offspring. We suggest that non-offspring nursing inlions occurs as a by-product of the females' communal defenseof their cubs against infanticide.     

ENDOSYMBIONTS OF DREISSENA POLYMORPHA IN IRELAND: EVIDENCE FOR THE INTRODUCTION OF ADULT MUSSELS

Although zebra mussels (Dreissena polymorpha) have invaded watersacross Europe for over 200 years, they colonized Ireland onlywithin the past decade. To test the hypothesis that Irelandwas colonized by adult D. polymorpha, we examined mussels fromdifferent sites along the Lower Shannon River system in Irelandfor the presence of host specific and generalist endosymbionts.Withinthe mantle cavity and/or associated with zebra mussel tissueswe found species specific-ciliates (Conchophthirus acuminatusand Ophryoglena hemophaga) and generalist symbionts (the ciliateAncistrumina limnica, nematodes, oligochaetes and chironomids).We found a significant difference in the prevalence of symbiontsamong sites, but all mussels at all sites harboured one specialistspecies C. acuminatus, and all of the mussels at three of thefour sites also had the second specialist, O. hemophaga. Thus,with the introduction of D. polymorpha into Ireland, at leasttwo additional species, their host-specific symbionts C. acuminatusand O. hemophaga, have also been introduced. The presence ofthese symbionts in Ireland supports the hypothesis that adultzebra mussels were introduced into Ireland, rather than larvalstages. This contrasts with the introduction of zebra musselsto North America, where adult zebra mussels are devoid of host-specificsymbionts. (Received 8 June 2005; accepted 7 November 2005)     

Grazing on colonial and filamentous, toxic and non-toxic cyanobacteria by the zebra mussel Dreissena polymorpha

Colony forming and toxic cyanobacteria form a problem in surfacewaters of shallow lakes, both for recreation and wildlife. Zebramussels, Dreissena polymorpha, have been employed to help torestore shallow lakes in the Netherlands, dominated by cyanobacteria,to their former clear state. Zebra mussels have been presentin these lakes since they were created in the 19th century bythe excavation of peat and are usually not considered to bean invasive species. Most grazing experiments using Dreissenahave been performed with uni-cellular phytoplankton laboratorystrains and information on grazing of larger phytoplankton taxahardly exists. To gain more insight in to whether D. polymorphais indeed able to decrease cyanobacteria in the phytoplankton,we therefore performed grazing experiments with zebra musselsand two species of cyanobacteria, that greatly differ in shape:colony forming strains of Microcystis aeruginosa and the filamentousspecies Planktothrix agardhii. For both species a toxic anda non-toxic strain was selected. We found that zebra musselscleared toxic Planktothrix at a higher rate than non-toxic Planktothrix,toxic or non-toxic Microcystis. Clearance rates between theother strains were not significantly different. Both phytoplanktonspecies, regardless of toxicity, size and shape, were foundin equal amounts (based on chlorophyll concentrations) in theexcreted products of the mussels (pseudofaeces). The resultsshow that zebra mussels are capable of removing colonial andfilamentous cyanobacteria from the water, regardless of whetherthe cyanobacteria are toxic or not. This implies that the musselsmay be used as a biofilter for the removal of harmful cyanobacterialblooms in shallow (Dutch) lakes where the mussels are alreadypresent and not a nuisance. Providing more suitable substratefor zebra mussel attachment may lead to appropriate mussel densitiescapable of filtering large quantities of cyanobacteria.     

Individual Foraging Strategies Reveal Niche Overlap between Endangered Galapagos Pinnipeds

Most competition studies between species are conducted from a population-level approach. Few studies have examined inter-specific competition in conjunction with intra-specific competition, with an individual-based approach. To our knowledge, none has been conducted on marine top predators. Sympatric Galapagos fur seals (Arctocephalus galapagoensis) and sea lions (Zalophus wollebaeki) share similar geographic habitats and potentially compete. We studied their foraging niche overlap at Cabo Douglas, Fernandina Island from simultaneously collected dive and movement data to examine spatial and temporal inter- and intra-specific competition. Sea lions exhibited 3 foraging strategies (shallow, intermediate and deep) indicating intra-specific competition. Fur seals exhibited one foraging strategy, diving predominantly at night, between 0–80 m depth and mostly at 19–22 h. Most sea lion dives also occurred at night (63%), between 0–40 m, within fur seals'' diving depth range. 34% of sea lions night dives occurred at 19–22 h, when fur seals dived the most, but most of them occurred at dawn and dusk, when fur seals exhibited the least amount of dives. Fur seals and sea lions foraging behavior overlapped at 19 and 21 h between 0–30 m depths. Sea lions from the deep diving strategy exhibited the greatest foraging overlap with fur seals, in time (19 h), depth during overlapping time (21–24 m), and foraging range (37.7%). Fur seals foraging range was larger. Cabo Douglas northwest coastal area, region of highest diving density, is a foraging “hot spot” for both species. Fur seals and sea lions foraging niche overlap occurred, but segregation also occurred; fur seals primarily dived at night, while sea lions exhibited night and day diving. Both species exploited depths and areas exclusive to their species. Niche breadth generally increases with environmental uncertainty and decreased productivity. Potential competition between these species could be greater during warmer periods when prey availability is reduced.     

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.     

Mountain lions on the prairie: habitat selection by recolonizing mountain lions at the edge of their range

Mountain lions (Puma concolor) have historically experienced large‐scale range contractions, but are beginning to recolonize portions of their former range. To reach potential suitable habitats in eastern North America, mountain lions need to move across the grassland and agriculture‐dominated habitats of the Great Plains, which are different from the forested areas associated with mountain lions in western North America. To inform restoration planning in this area, it is important to understand differences in mountain lion habitat selection in this “nontraditional” grassland habitat. We tracked GPS‐collared mountain lions in the Northern Great Plains of Montana and identified movement states (localized or exploratory) using behavioral change point analysis and net‐squared displacement. We compared habitat selection between the different states using step‐selection functions that included several environmental covariates. Similar to elsewhere throughout their range, across both movement states, mountain lions tended to select forested environments that were farther from human development. In contrast to more traditionally occupied mountainous regions, mountain lions in the Great Plains selected areas of lower elevations. They selected areas both near and far from water, but avoided riparian areas and selected more rugged environments when in exploratory movement states. This suggests that mountain lions in the Northern Great Plains are utilizing river corridors, particularly those with rough or broken topography during exploratory phases. To enhance future recolonization and connectivity of mountain lions to the east of our study area, we encourage managers to maintain and restore forest fragments along river corridors in the Great Plains.     

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.