Distributional niche of relatively rare sable antelope in a South African savanna: habitat versus biotic relationships

The geographic distribution of a species is governed by climatic conditions, topography, resources and habitat structure determining the fundamental niche, while the local distribution expressed via home range occupation may be compressed by biotic interactions with competitors and predators, restricting the realised niche. Biotic influences could be especially important for relatively rare species. We investigated how rainfall, geology, land type and abundance of other ungulate species serving as competitors or prey for predators contributed to the patchy distribution of sable antelope herds within Kruger National Park. Data were provided by annual aerial surveys of ungulate populations conducted between 1978 and 1988. Sable herds were more commonly present on granitic and sandstone substrates than on more fertile basalt. They occurred both in the moist south‐west and dry north of the park. They were most abundant in sour bushveld and mopane savanna woodland, and mostly absent from knob thorn‐marula parkland. The presence of sable was negatively associated with high concentrations of impala and wildebeest, less consistently related to the abundance of zebra, and positively associated with the occurrence of buffalo herds. Best supported models included the separate effects of the most abundant grazers along with land type. Interspecific relationships seemed more consistent with vulnerability to predation as the underlying mechanism restricting the distribution of sable herds than with competitive displacement. Sable favoured land types distinct from those where wildebeest, the most preferred prey of lions, and impala, numerically the most important resident prey species, were most abundant. Hence the risk of predation, associated with habitat conditions where abundant prey species are most concentrated, can exert an overriding influence on the distribution of rarer species in terms of their home range occupation.     

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.     

Changing vulnerability to predation related to season and sex in an African ungulate assemblage

The consequences of predation for prey population dynamics depend on the extent to which this mortality is predisposed by malnutrition or senescence, or additive in the sense that animals that would otherwise not have died at that time were killed. In places lacking effective predators, few adult ungulates die during the summer or wet season months when food is plentifully available. Hence the seasonal distribution of predator kills as well as the age and sex classes of the prey indicates the extent to which malnutrition contributes to mortality as well as other influences on vulnerability. Using records of animal deaths assembled over 35 years in South Africa's Kruger National Park, these patterns were investigated for 12 ungulate species forming the prey of lions, and for three other large predators with respect to one prey species. Buffalo, kudu and giraffe were more strongly represented in kills made during the late dry season, while wildebeest and zebra made relatively greater contributions during the wet season. Impala, waterbuck, warthog and rarer antelope species became more prominent in kills during transitional periods between seasons. Five prey species showed an elevation in representation of males in lion kills during the mating season, as well as impala for all predator species. Females were more prominently represented in kills during the time of late gestation and parturition for three prey species. Hence reproductive activities as well as changing vegetation cover and food resources affected vulnerability to predation. Shifts in susceptibility to predation over the seasonal cycle corresponded with rainfall‐related variation in the annual representation of these ungulate species in lion kills. The availability of vulnerable prey species, age and sex classes at different stages of the seasonal cycle helps maintain a high abundance of lions. These factors contribute to the strong additive impact that predation has had on the abundance of some of these ungulate populations.     

The pattern of lion predation in Nairobi Park

The feeding habits of lions in Nairobi National Park have been observed between 1968 and 1972. Nairobi Park is briefly described and the herbivore and predator populations are discussed. The relationship between the changing status of the prey population for the period 1968-72 and the feeding habits of the lions is examined and compared with data from earlier years. Selection ratio (preference ratio) for three of the most important prey species is more closely adjusted to the abundance of these species in the total ungulate population than was the case prior to 1968. Only warthog and eland form an exception. Predation on wildebeest shows the most marked decrease although this species has been increasing during the last 5 years. Wildebeest, kongoni, zebra and warthog have provided 80% of total kills throughout the study on the basis of numbers killed. On this basis, warthog has the highest preference rating. However, the largest contribution to the lions' diet expressed in kg is by kongoni, followed by zebra, wildebeest, eland and warthog, in that order. Kongoni and wildebeest contribute most during the first half of the year and zebra and eland during the last 6 months. Selection for males is significant in zebra and eland and highly significant in kongoni, whilst only in zebra is selection for juveniles evident.     

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.     

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.     

Large herbivore assemblages in a changing climate: incorporating water dependence and thermoregulation

The coexistence of different species of large herbivores (ungulates) in grasslands and savannas has fascinated ecologists for decades. However, changes in climate, land‐use and trophic structure of ecosystems increasingly jeopardise the persistence of such diverse assemblages. Body size has been used successfully to explain ungulate niche differentiation with regard to food requirements and predation sensitivity. But this single trait axis insufficiently captures interspecific differences in water requirements and thermoregulatory capacity and thus sensitivity to climate change. Here, we develop a two‐dimensional trait space of body size and minimum dung moisture content that characterises the combined food and water requirements of large herbivores. From this, we predict that increased spatial homogeneity in water availability in drylands reduces the number of ungulate species that will coexist. But we also predict that extreme droughts will cause the larger, water‐dependent grazers as wildebeest, zebra and buffalo–dominant species in savanna ecosystems – to be replaced by smaller, less water‐dependent species. Subsequently, we explore how other constraints such as predation risk and thermoregulation are connected to this two‐dimensional framework. Our novel framework integrates multiple simultaneous stressors for herbivores and yields an extensive set of testable hypotheses about the expected changes in large herbivore community composition following climate change.     

Modelling ungulate dependence on higher quality forage under large trees in African savannahs

In African savannahs, large trees improve grass quality, particularly in dry and nutrient poor areas. Enhanced below-canopy grass nutrients, such as nitrogen and phosphorus contents should therefore attract and benefit grazers. To predict whether ungulates really need these forage quality islands we focused on four grazer species, i.e., zebra, buffalo, wildebeest, and warthog, differing in body size and digestive system. We confronted literature estimations of their feeding requirements with forage availability and quality, observed in three South African savannah systems, through linear modelling. The model predicted the proportion of below-canopy grass that grazers should include in their diet to meet their nutritional requirements.During the wet season, the model predicted that all animals could satisfy their daily nutrient requirements when feeding on a combination of below- and outside-canopy grasses. However, wildebeest, having relatively high nutrient demands, could meet their nutrient requirements only by feeding almost exclusively below canopies.During the dry season, all animals could gain almost twice as much digestible protein when feeding on below – compared to outside-canopy forage. Nonetheless, only warthogs could satisfy their nutrient requirements – when feeding almost exclusively on below-canopy grasses. The other ungulate species could not meet their phosphorus demands by feeding at either site without exceeding their maximum fibre intake, indicating the unfavourable conditions during the dry season.We conclude that grazing ungulates, particularly warthog, zebra, and buffalo, actually depend on the available below-canopy grass resources. Our model therefore helps to quantify the importance of higher quality forage patches beneath savannah trees. The composition of grazer communities depending on below-canopy grasses can be anticipated if grazer food requirements and the abundance of large trees in savannahs are known. The model suggests that the conservation of large single-standing trees in savannahs is crucial for maintenance of locally grazing herbivores.     

Migratory herds of wildebeests and zebras indirectly affect calf survival of giraffes

相似文献   

Ecological and anthropogenic effects on the density of migratory and resident ungulates in a human‐inhabited protected area

Understanding the influence of environmental conditions and people on ungulate density and distribution is of key importance for conservation. We evaluated the effects of ecological and anthropogenic factors on the density of migratory wildebeest and zebra and resident oribi in Zambia's Liuwa Plain National Park where human settlements were present. We conducted transect surveys from 2010 to 2013 using distance sampling methods and then developed a set of 38 candidate models to describe results and predict density. Models included the effects of variables in three classes: environmental (year, season, vegetation, predominant grass height, burn, water presence), predation risk (hyaena density) and anthropogenic (distance to park boundary and settlements). Densities ranged from 6.2 to 60.8 individuals km^?2 for wildebeest, 1.1 to 14.5 individuals km^?2 for oribi and 1.8 to 8.1 individuals km^?2 for zebra. The most complex models were strongly supported for all three species. The magnitude and sign of variable effects differed among species, indicating that local densities of wildebeest, oribi and zebra are affected by a complex set of anthropogenic and ecological factors. Results reveal resource partitioning among ungulate species and indicate that predation risk and proximity to humans affect ungulate distributions with implications for managing migrations in the Greater Liuwa Ecosystem.     

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).     

Demography of plains zebras (Equus quagga) under heavy predation

In natural ecosystems, ungulate densities show strong temporal variations. The ecological processes driving these fluctuations are complex: food limitation and predation are both important and can interact. Survival rates are central to this debate, but data are sparse for tropical ecosystems. Here, we estimate age- and sex-specific survival rates for plains zebra in Hwange National Park, a nutrient-poor savanna with a high predator–prey ratio. We estimated survival from a detailed Capture-Mark-Recapture (CMR) monitoring based on 248 individual life histories, for the first time in an African grazer. We controlled for variations in detection probabilities among adult females, which resulted from their social structure. As expected, annual survival was low during the first year (0.441); increased in yearlings (0.560) and peaked at 0.795 and 0.847 in adult males and females respectively. The survival of adult females was lower during the dry season, which probably resulted from higher predation due to predictable movements of zebras to waterholes. Survival at all ages was low compared to ungulates without predators. The demographic model we constructed showed a declining trend (λ = 0.94), which was consistent with the data from road counts ( = 0.92). Life Table Response Experiment (LTRE) analyses using the Serengeti and Kruger populations as references showed that the main cause of this declining trend in the Hwange population was low survival in yearling and adult females; low foal survival also contributed. In this ecosystem, predation is likely to be the main ecological process causing low survival, and therefore a decline in the zebra population.     

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.     

Dynamic spatial partitioning and coexistence among tall grass grazers in an African savanna

Competitive relationships among mobile animals may be expressed through dynamically changing spatial relationships over different time frames. Less common species that are apparently inferior competitors may be able to coexist with more abundant species by concentrating in regions of the landscape little utilized by the former at spatio‐temporal scales from annual or seasonal ranges to the specific foraging localities exploited at different stages of the annual cycle. Spatial relationships may be influenced further by dependencies on other resources, predation risks and facilitatory interactions under certain conditions. Our study aimed to determine whether competition with more abundant zebra and buffalo restricted the abundance of sable antelope in a region where these three tall‐grass grazers overlapped in their herd distributions. We tracked the simultaneous movements of animals representing herds of these species over two dry seasons and one wet season using GPS‐GSM collars, and estimated seasonal or monthly range extents and their overlap. We also compared daily separation distances between these animals against the null pattern expected if their movements had been independent, and assessed how prior grazing by buffalo influenced the subsequent use of these localities by sable. The range of the sable herd was mostly separated from the seasonal range of the buffalo herd during the late dry season of 2006 and throughout the dry season of 2007. Seasonal home ranges of zebra herds overlapped partially with the range of the sable herd during most of the year. Even during times when their ranges overlapped, sable were rarely recorded within <1 km of the buffalo herd. Prior grazing by buffalo beyond a threshold level inhibited later use of these localities by sable, but the sable were nevertheless able to exploit places that were little utilized by buffalo at that time. Sable were less able to evade overlap with the small, mobile zebra herds, and hence more vulnerable to competitive exclusion by zebra than by buffalo. Our findings demonstrate how less abundant species can restrict competition from more abundant competitors through dynamic spatial partitioning in regions where their home ranges overlap.     

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.     

Biotic resistance on the increase: native predators structure invasive zebra mussel populations

1. Abundant native predators, parasites and pathogens that switch to consuming a hyper‐successful exotic species may be able to control the invasive population. Native predators may, however, need time to adapt to feed effectively on an exotic resource. In this case, mortality on an exotic population from native predators could increase over time even without a numerical increase in the predator population. 2. We measured mortality of zebra mussels (Dreissena polymorpha) in the Hudson River both in controls open to predation and in exclosures that excluded large predators to estimate mortality of zebra mussels from large predators and other causes. 3. We found that predation by the blue crab (Callinectes sapidus), and perhaps other predators, causes high mortality on zebra mussels in the Hudson River estuary. This predation apparently led to increased mortality and altered population structure in the invader over time. 4. Long‐term data from the Hudson River suggest that components of the invaded ecosystem, like rotifers, are recovering through predator‐caused release from zebra mussel grazing. Increased mortality on hyper‐successful exotic populations over time may be a common phenomenon with both ecological and management implications.     

Predation on invasive zebra mussel, Dreissena polymorpha, by pumpkinseed sunfish, rusty crayfish, and round goby

The enemy release hypothesis states that invasive species are successful in their new environment because native species are not adapted to utilize the invasive. If true for predators, native predators should have lower feeding rates on the invasive species than a predator from the native range of the invasive species. We tested this hypothesis for zebra mussel (Dreissena polymorpha) by comparing handling time and predation rate on zebra mussels in the laboratory by two North American species (pumpkinseed, Lepomis gibbosus, and rusty crayfish, Orconectes rusticus) and one predator with a long evolutionary history with zebra mussels (round goby, Neogobius melanostomus). Handling time per mussel (7 mm shell length) ranged from 25 to >70 s for the three predator species. Feeding rates on attached zebra mussels were higher for round goby than the two native predators. Medium and large gobies consumed 50–67 zebra mussels attached to stones in 24 h, whereas pumpkinseed and rusty crayfish consumed <11. This supports the hypothesis that the rapid spread of zebra mussels in North America was facilitated by low predation rates from the existing native predators. At these predation rates and realistic goby abundance estimates, round goby could affect zebra mussel abundance in some lakes.     

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.     

Group Dynamics of Zebra and Wildebeest in a Woodland Savanna: Effects of Predation Risk and Habitat Density

Background

Group dynamics of gregarious ungulates in the grasslands of the African savanna have been well studied, but the trade-offs that affect grouping of these ungulates in woodland habitats or dense vegetation are less well understood. We examined the landscape-level distribution of groups of blue wildebeest, Connochaetes taurinus, and Burchell''s zebra, Equus burchelli, in a predominantly woodland area (Karongwe Game Reserve, South Africa; KGR) to test the hypothesis that group dynamics are a function of minimizing predation risk from their primary predator, lion, Panthera leo.

Methodology/Principal Findings

Using generalized linear models, we examined the relative importance of habitat type (differing in vegetation density), probability of encountering lion (based on utilization distribution of all individual lions in the reserve), and season in predicting group size and composition. We found that only in open scrub habitat, group size for both ungulate species increased with the probability of encountering lion. Group composition differed between the two species and was driven by habitat selection as well as predation risk. For both species, composition of groups was, however, dominated by males in open scrub habitats, irrespective of the probability of encountering lion.

Conclusions/Significance

Distribution patterns of wildebeest and zebra groups at the landscape level directly support the theoretical and empirical evidence from a range of taxa predicting that grouping is favored in open habitats and when predation risk is high. Group composition reflected species-specific social, physiological and foraging constraints, as well as the importance of predation risk. Avoidance of high resource open scrub habitat by females can lead to loss of foraging opportunities, which can be particularly costly in areas such as KGR, where this resource is limited. Thus, landscape-level grouping dynamics are species specific and particular to the composition of the group, arising from a tradeoff between maximizing resource selection and minimizing predation risk.     

Predation, sex ratios, and male competition in equids (Mammalia: Perissodactyla)

Existing data indicate that a greater preponderance of adult females rather than adult males occurs in most species of mammals. The hypothesis that such differences arise as a result of intermale reproductive competition for females (and not predation) was examined in the Equidae by comparing populations of horses ( Equus caballus ), asses ( E. asinus ), and two species of zebras ( E. zebra and E. burchelli) in predator-free, predator-rich and insular ecosystems.
Evidence is presented that: (1) sex differences in adult mortality occur; (2) they relate to the type and intensity of natural predation; and (3) asymmetries in sex ratios are most often explicable in terms of intermale reproductive competition. Exceptions are discussed and they are complicated by numerous proximate factors.