Predation strongly limits demography of a keystone migratory herbivore in a recovering transfrontier ecosystem

Large herbivore migrations are imperiled globally; however the factors limiting a population across its migratory range are typically poorly understood. Zambia's Greater Liuwa Ecosystem (GLE) contains one of the largest remaining blue wildebeest (Connochaetes taurinus taurinus) migrations, yet the population structure, vital rates, and limiting factors are virtually unknown. We conducted a long-term demographic study of GLE wildebeest from 2012 to 2019 of 107 collared adult females and their calves, 7352 herd observations, 12 aerial population surveys, and concurrent carnivore studies. We applied methods of vital rate estimation and survival analysis within a Bayesian estimation framework. From herd composition observations, we estimated rates of fecundity, first-year survival, and recruitment as 68%, 56%, and 38% respectively, with pronounced interannual variation. Similar rates were estimated from calf-detections with collared cows. Adult survival rates declined steadily from 91% at age 2 years to 61% at age 10 years thereafter dropping more sharply to 2% at age 16 years. Predation, particularly by spotted hyena, was the predominant cause of death for all wildebeest ages and focused on older animals. Starvation only accounted for 0.8% of all unbiased known natural causes of death. Mortality risk differed substantially between wet and dry season ranges, reflecting strong spatio-temporal differences in habitat and predator densities. There was substantial evidence that mortality risk to adults was 27% higher in the wet season, and strong evidence that it was 45% higher in the migratory range where predator density was highest. The estimated vital rates were internally consistent, predicting a stable population trajectory consistent with aerial estimates. From essentially zero knowledge of GLE wildebeest dynamics, this work provides vital rates, age structure, limiting factors, and a plausible mechanism for the migratory tendency, and a robust model-based foundation to evaluate the effects of potential restrictions in migratory range, climate change, predator-prey dynamics, and poaching

Habitat shifts in response to predation risk are constrained by competition within a grazing guild

Predators can affect prey not only by killing them, but also by causing them to alter their behavior, including patterns of habitat selection. Prey can reduce the risk of predation by moving to habitats where predators are less likely to detect them, less likely to attack, or less likely to succeed. The interaction of such responses to risk with other ecological processes remains relatively unstudied, but in some cases, changes in habitat use to avoid predation may be constrained by competition: larger, dominant competitors should respond freely to predation risk, but the responses of smaller, subordinate competitors may be constrained by the responses of dominant competitors. For large grazing herbivores, an alternative hypothesis proposes that smaller prey species are vulnerable to more predators, and thus should respond more strongly to predation risk. Here, we tested these two hypotheses with 775 observations of habitat selection by four species of obligate grazers (zebra, wildebeest, puku and oribi) in the immediate presence or absence of four large carnivores (lion, spotted hyena, African wild dog and cheetah) in three ecosystems (Greater Liuwa, Greater Kafue and Luangwa Valley). Patterns of predation within this set were described by observation of 1,105 kills. Our results support the hypothesis that responses to predation risk are strongest for larger, dominant competitors. Even though zebras were killed least often, they showed the strongest shift into cover when carnivores were present. Wildebeest, puku and oribi showed weaker habitat shifts, even though they were more frequently killed. These patterns remained consistent in models that controlled for differences in the hunting mode of the predator (stalking, coursing, or intermediate) and for differences among ecosystems. There was no evidence that smaller species were subject to predation by a broader set of predators. Instead, smaller prey were killed often by smaller predators, and larger prey were killed often by larger predators. Broadly, our results show that responses to predation risk interact with interspecific competition. Accounting for such interactions should help to explain the considerable variation in the strength of responses to predation risk that has been observed

Using dogs to find cats: detection dogs as a survey method for wide-ranging cheetah

Rapid global large carnivore declines make evaluations of remaining populations critical. Yet landscape-scale evaluations of presence, abundance and distribution are difficult, as many species are wide-ranging, occur only at low densities and are elusive. Insufficient information-gathering tools for many large carnivore species compounds these challenges. Specially trained detection dogs have demonstrated effectiveness for carnivore surveys, but are untested on extremely sparse, wide-ranging species, such as cheetah Acinonyx jubatus. In this study, we conducted the first rigorous cheetah survey using detection dogs in a key transboundary area in the remote Liuwa–Mussuma Transfrontier Conservation Area (TFCA) in Western Zambia. We proposed to (1) evaluate the effectiveness of detection dog versus spoor surveys in detecting cheetah presence; (2) extract and analyze DNA from scat samples to estimate minimum population size and genetic effective population size; (3) determine the extent of cheetah occurrence in the unprotected transboundary corridor. Two detection dog teams surveyed 2432 km2 containing 74 randomly located transects in the transfrontier area. Twenty-seven cheetah scats were detected and confirmed by genetic analysis, while no cheetah spoor was detected, clearly demonstrating the superiority of detection dogs in detecting cheetah presence. Combining scat samples with opportunistically collected samples, we estimated 17–19 cheetahs, an effective population size of 8–14 and a density of 5.9–6.6 per 1000 km2. Cheetah utilized key transfrontier areas outside of the national park; however, because utilization appears low, improved connectivity and protection for these areas are critical. Approximately one third of Africa's estimated cheetah resides in protected areas, with 87% in transboundary areas. Our study demonstrates the efficacy of detection dog survey methods in providing information on cheetah across large landscapes. It will have particular value in areas where other survey means may be impossible, such as TFCAs, where size, remoteness and lack of accessibility often make traditional survey methods difficult or cost prohibitive

Cultural and health beliefs of pregnant women in Zambia regarding pregnancy and child birth

Background: Health beliefs related to pregnancy and childbirth exist in various cultures globally. Healthcare practitioners need to be aware of these beliefs so as to contextualise theirpractice in their communities. Objectives: To explore the health beliefs regarding pregnancy and childbirth of womenattending the antenatal clinic at Chawama Health Center in Lusaka Zambia. Method: This was a descriptive, cross-sectional survey of women attending antenatal care(n = 294) who were selected by systematic sampling. A researcher-administered questionnairewas used for data collection. Results: Results indicated that women attending antenatal care at Chawama Clinic held certain beliefs relating to diet, behaviour and the use of medicinal herbs during pregnancy and post-delivery. The main beliefs on diet related to a balanced diet, eating of eggs, okra, bones, offal, sugar cane, alcohol consumption and salt intake. The main beliefs on behaviour related to commencement of antenatal care, daily activities, quarrels, bad rituals, infidelity and the use of condoms during pregnancy. The main beliefs on the use of medicinal herbs were on their use to expedite the delivery process, to assist in difficult deliveries and for body cleansing following a miscarriage. Conclusion: Women attending antenatal care at the Chawama Clinic hold a number of beliefs regarding pregnancy and childbirth. Those beliefs that are of benefit to the patients should be encouraged with scientific explanations, whilst those posing a health risk should be discouraged respectfully

Spatial and temporal avoidance of risk within a large carnivore guild

Abstract Within a large carnivore guild, subordinate competitors (African wild dog, Lycaon pictus, and cheetah, Acinonyx jubatus) might reduce the limiting effects of dominant competitors (lion, Panthera leo, and spotted hyena, Crocuta crocuta) by avoiding them in space, in time, or through patterns of prey selection. Understanding how these competitors cope with one other can inform strategies for their conservation. We tested how mechanisms of niche partitioning promote coexistence by quantifying patterns of prey selection and the use of space and time by all members of the large carnivore guild within Liuwa Plain National Park in western Zambia. Lions and hyenas specialized on wildebeest, whereas wild dogs and cheetahs selected broader diets including smaller and less abundant prey. Spatially, cheetahs showed no detectable avoidance of areas heavily used by dominant competitors, but wild dogs avoided areas heavily used by lions. Temporally, the proportion of kills by lions and hyenas did not detectably differ across four time periods (day, crepuscular, early night, and late night), but wild dogs and especially cheetahs concentrated on time windows that avoided nighttime hunting by lions and hyenas. Our results provide new insight into the conditions under which partitioning may not allow for coexistence for one subordinate species, the African wild dog, while it does for cheetah. Because of differences in responses to dominant competitors, African wild dogs may be more prone to competitive exclusion (local extirpation), particularly in open, uniform ecosystems with simple (often wildebeest dominated) prey communities, where spatial avoidance is difficult

Risky times and risky places interact to affect prey behaviour.

Both short-term and long-term variation in predation risk can affect the behaviour of prey, thus affecting growth, reproduction, survival and population dynamics. Inferences about the strength of such 'risk effects' in the wild have been limited by a lack of studies that relate antipredator responses to the magnitude of direct predation, measure responses of prey to risk from complete predator guilds, and quantify risk in more than one way. Here, we quantify behavioural responses of a complete ungulate prey guild to long-term and short-term variation in risks from all of the large predators in Liuwa Plain National Park, with known patterns of direct predation. Our analysis allows the first direct test for interaction between responses to long-term and short-term risk in the wild, and reveals that prey vigilance responds strongly to locations with high long-term risk when short-term risk is high, but not when short-term risk is low. This result has broad ramifications for the design and interpretation of field studies of antipredator behaviour, its costs and its consequences for population dynamics

Risky times and risky places interact to affect prey behaviour.

Both short-term and long-term variation in predation risk can affect the behaviour of prey, thus affecting growth, reproduction, survival and population dynamics. Inferences about the strength of such 'risk effects' in the wild have been limited by a lack of studies that relate antipredator responses to the magnitude of direct predation, measure responses of prey to risk from complete predator guilds, and quantify risk in more than one way. Here, we quantify behavioural responses of a complete ungulate prey guild to long-term and short-term variation in risks from all of the large predators in Liuwa Plain National Park, with known patterns of direct predation. Our analysis allows the first direct test for interaction between responses to long-term and short-term risk in the wild, and reveals that prey vigilance responds strongly to locations with high long-term risk when short-term risk is high, but not when short-term risk is low. This result has broad ramifications for the design and interpretation of field studies of antipredator behaviour, its costs and its consequences for population dynamics

Response of wildebeest (Connochaetes taurinus) movements to spatial variation in long term risks from a complete predator guild

Many studies have shown that behavioral responses to the risk posed by predators can carry costs for prey by reducing fecundity or survival, with consequent effects on population dynamics. Responses to risk include increased vigilance and reduced foraging, movement to safe habitats, increases or decreases in group size, and changes in patterns of movement. While we know that prey can detect and respond to both long term (LT) and short term (ST) variation in risk, field studies have only recently begun to consider how these responses might differ. Here, we hypothesize that prey movement patterns should respond differently to cues of LT and ST variation in risk. Specifically, cues of elevated LT risk might lead to decreased movement to improve the assessment of ST risk, while elevated ST risk might favor increased movement to reduce the proximity or duration of risks that are already assessed to be acute. We further hypothesize that decreases in movement are likely to be a general response to LT risk, while responses to ST risk are likely to vary in a manner that depends on the type of predator. In Liuwa Plain National Park, we found that wildebeest movements responded to the local intensity of predator use (LT risk), after controlling for other seasonal, diurnal and bottom-up effects. Speed decreased considerably and turning angles increased considerably, combining to markedly decrease linear movements. In contrast, immediate encounters with predators (ST risk) typically provoked fast, linear flight, and this effect was stronger for encounters with coursing predators. The effect of long term risk was to cause wildebeest to move more slowly and less linearly, i.e. to slow down and turn around, as part of a suite of behavioral responses, which also includes increased vigilance, that promote cautious assessment of ST risks when in locations with high levels of LT risk. This result has broad implications of understanding the influence of predation risk on foraging patterns of ungulates as this relationship is much more complex than simple avoidance of areas of ‘high risk’

Response of wildebeest (Connochaetes taurinus) movements to spatial variation in long term risks from a complete predator guild

Many studies have shown that behavioral responses to the risk posed by predators can carry costs for prey by reducing fecundity or survival, with consequent effects on population dynamics. Responses to risk include increased vigilance and reduced foraging, movement to safe habitats, increases or decreases in group size, and changes in patterns of movement. While we know that prey can detect and respond to both long term (LT) and short term (ST) variation in risk, field studies have only recently begun to consider how these responses might differ. Here, we hypothesize that prey movement patterns should respond differently to cues of LT and ST variation in risk. Specifically, cues of elevated LT risk might lead to decreased movement to improve the assessment of ST risk, while elevated ST risk might favor increased movement to reduce the proximity or duration of risks that are already assessed to be acute. We further hypothesize that decreases in movement are likely to be a general response to LT risk, while responses to ST risk are likely to vary in a manner that depends on the type of predator. In Liuwa Plain National Park, we found that wildebeest movements responded to the local intensity of predator use (LT risk), after controlling for other seasonal, diurnal and bottom-up effects. Speed decreased considerably and turning angles increased considerably, combining to markedly decrease linear movements. In contrast, immediate encounters with predators (ST risk) typically provoked fast, linear flight, and this effect was stronger for encounters with coursing predators. The effect of long term risk was to cause wildebeest to move more slowly and less linearly, i.e. to slow down and turn around, as part of a suite of behavioral responses, which also includes increased vigilance, that promote cautious assessment of ST risks when in locations with high levels of LT risk. This result has broad implications of understanding the influence of predation risk on foraging patterns of ungulates as this relationship is much more complex than simple avoidance of areas of ‘high risk’