Browsing by Author "Droge, Egil"

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Assessing the Performance of Index Calibration Survey Methods to Monitor Populations of Wide‐ranging Low‐density Carnivores
(2020-03) Droge, Egil; Creel, Scott; Becker, Matthew S.; Loveridge, Andrew J.; Sousa, Lara L.; Macdonald, David W.
Apex carnivores are wide‐ranging, low‐density, hard to detect, and declining throughout most of their range, making population monitoring both critical and challenging. Rapid and inexpensive index calibration survey (ICS) methods have been developed to monitor large African carnivores. ICS methods assume constant detection probability and a predictable relationship between the index and the actual population of interest. The precision and utility of the resulting estimates from ICS methods have been questioned. We assessed the performance of one ICS method for large carnivores—track counts—with data from two long‐term studies of African lion populations. We conducted Monte Carlo simulation of intersections between transects (road segments) and lion movement paths (from GPS collar data) at varying survey intensities. Then, using the track count method we estimated population size and its confidence limits. We found that estimates either overstate precision or are too imprecise to be meaningful. Overstated precision stemmed from discarding the variance from population estimates when developing the method and from treating the conversion from tracks counts to population density as a back‐transformation, rather than applying the equation for the variance of a linear function. To effectively assess the status of species, the IUCN has set guidelines, and these should be integrated in survey designs. We propose reporting the half relative confidence interval width (HRCIW) as an easily calculable and interpretable measure of precision. We show that track counts do not adhere to IUCN criteria, and we argue that ICS methods for wide‐ranging low‐density species are unlikely to meet those criteria. Established, intensive methods lead to precise estimates, but some new approaches, like short, intensive, (spatial) capture–mark–recapture (CMR/SECR) studies, aided by camera trapping and/or genetic identification of individuals, hold promise. A handbook of best practices in monitoring populations of apex carnivores is strongly recommended.
Detecting declines of apex carnivores and evaluating their causes: An example with Zambian lions
(Elsevier, 2014) Rosenblatt, Elias G.; Becker, Matthew S.; Creel, Scott; Droge, Egil; Mweetwa, Thandiwe; Schuette, Paul; Watson, Fred; Merkle, Johnathan; Mwape, Henry
Large carnivores are in rapid global decline, with a broad array of consequences for the ecosystems they inhabit. To efﬁciently detect and address these declines requires unbiased and precise demographic data. Unfortunately, the characteristics that make large carnivores extinction-prone also pose serious challenges to obtaining these data. Rapid survey methods exist, but provide only relative measures of abundance, cannot detect declines before they become large, and provide little or no information about the causes of decline. African lions (Panthera leo) are declining throughout their range, making accurate monitoring of remaining populations urgent. We provide statistically rigorous estimates of population size, trends, survival rate and age–sex structure from Zambia’s South Luangwa lion population from 2008 to 2012, just prior to cessation of hunting in 2013. Mark-recapture models ﬁt to data from intensive monitoring of 210 individual lions in 18 prides and 14 male coalitions indicated a declining population, low recruitment, low sub-adult and adult male survival, depletion of adult males, and a senescing adult female population. Trophy hunting was the leading cause of death, with 46 males harvested. Based on these data we recommend continuing the hunting ban at least to 2016 to allow recovery, with substan-tially reduced quotas, age-limits, and effective trophy monitoring mandated thereafter should hunting resume. Similar data from intensive monitoring of key Zambian lion populations is required to evaluate effects of the hunting ban and provide management guidance. Effectively integrating intensive long-term monitoring and rapid survey methods should be a priority for future management and monitoring of carnivore species.
Foraging investment in a long-lived herbivore and vulnerability to coursing and stalking predators
(2018-10) Christianson, David A.; Becker, Matthew S.; Brennan, Angela; Creel, Scott; Droge, Egil; M'soka, Jassiel; Mukula, Teddy; Schuette, Paul; Smit, Daan; Watson, Fred
Allocating resources to growth and reproduction requires grazers to invest time in foraging, but foraging promotes dental senescence and constrains expression of proactive antipredator behaviors such as vigilance. We explored the relationship between carnivore prey selection and prey foraging effort using incisors collected from the kills of coursing and stalking carnivores. We predicted that prey investing less effort in foraging would be killed more frequently by coursers, predators that often exploit physical deficiencies. However, such prey could expect delayed dental senescence. We predicted that individuals investing more effort in foraging would be killed more frequently by stalkers, predators that often exploit behavioral vulnerabilities. Further these prey could expect earlier dental senescence. We tested these predictions by comparing variation in age-corrected tooth wear, a proxy of cumulative foraging effort, in adult (3.4-11.9 years) wildebeest killed by coursing and stalking carnivores. Predator type was a strong predictor of age-corrected tooth wear within each gender. We found greater foraging effort and earlier expected dental senescence, equivalent to 2.6 additional years of foraging, in female wildebeest killed by stalkers than in females killed by coursers. However, male wildebeest showed the opposite pattern with the equivalent of 2.4 years of additional tooth wear in males killed by coursers as compared to those killed by stalkers. Sex-specific variation in the effects of foraging effort on vulnerability was unexpected and suggests that behavioral and physical aspects of vulnerability may not be subject to the same selective pressures across genders in multipredator landscapes.
Predation strongly limits demography of a keystone migratory herbivore in a recovering transfrontier ecosystem
(Wiley, 2022-10) Watson, Fred; Becker, Matthew S.; Smit, Daan; Droge, Egil; Mukula, Teddy; Martens, Sandra; Mwaba, Shadrach; Christianson, David; Creel, Scott; Brennan, Angela; M'soka, Jassiel; Gaylard, Angela; Simukonda, Chuma; Nyirenda, Moses; Mayani, Bridget
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.
Quantifying lion (Panthera leo) demographic response following a three-year moratorium on trophy hunting.
(2018-05) Mweetwa, Thandiwe; Christianson, David A.; Becker, Matthew S.; Creel, Scott; Rosenblatt, Elias G.; Merkle, Johnathan; Droge, Egil; Mwape, Henry; Masonde, Jones; Simpamba, Twakundine
Factors that limit African lion populations are manifold and well-recognized, but their relative demographic effects remain poorly understood, particularly trophy hunting near protected areas. We identified and monitored 386 individual lions within and around South Luangwa National Park, Zambia, for five years (2008-2012) with trophy hunting and for three additional years (2013-2015) during a hunting moratorium. We used these data with mark-resight models to estimate the effects of hunting on lion survival, recruitment, and abundance. The best survival models, accounting for imperfect detection, revealed strong positive effects of the moratorium, with survival increasing by 17.1 and 14.0 percentage points in subadult and adult males, respectively. Smaller effects on adult female survival and positive effects on cub survival were also detected. The sex-ratio of cubs shifted from unbiased during trophy-hunting to female-biased during the moratorium. Closed mark-recapture models revealed a large increase in lion abundance during the hunting moratorium, from 116 lions in 2012 immediately preceding the moratorium to 209 lions in the last year of the moratorium. More cubs were produced each year of the moratorium than in any year with trophy hunting. Lion demographics shifted from a male-depleted population consisting mostly of adult (â‰¥4 years) females to a younger population with more (>29%) adult males. These data show that the three-year moratorium was effective at growing the Luangwa lion population and increasing the number of adult males. The results suggest that moratoria may be an effective tool for improving the sustainability of lion trophy hunting, particularly where systematic monitoring, conservative quotas, and age-based harvesting are difficult to enforce.
Spatial and temporal avoidance of risk within a large carnivore guild
(2017-01) Droge, Egil; Creel, Scott; Becker, Matthew S.; M'soka, Jassiel
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.
Spatial Risk Effects From Lions Compound Impacts of Prey Depletion on African Wild Dogs
(Wiley, 2024-10) Goodheart, Ben; Creel, Scott; Schuette, Paul; Droge, Egil; Becker, Justine A.; Banda, Kambwiri; Kusler, Anna; Matsushima, Stephi; Banda, Kachama; Kabwe, Ruth; Donald, Will; Reyes de Merkle, Johnathan; Kaluka, Adrian; Chifunte, Clive; Becker, Matthew S.
Prey depletion threatens many carnivore species across the world and can especially threaten low-density subordinate competitors, particularly if subordinates are limited to low densities by their dominant competitors. Understanding the mechanisms that drive responses of carnivore density to prey depletion is not only crucial for conservation but also elucidates the balance between top-down and bottom-up limitations within the large carnivore guild. To avoid predation, competitively subordinate African wild dogs typically avoid their dominant competitors (lions) and the prey rich areas they are associated with, but no prior research has tested whether this pattern persists in ecosystems with anthropogenically-reduced prey density, and reduced lion density as a result. We used spatial data from wild dogs and lions in the prey-depleted Greater Kafue Ecosystem to test if wild dogs continue to avoid lions (despite their low density), and consequently avoid habitats with higher densities of their dominant prey species. We found that although lion density is 3X lower than comparable ecosystems, wild dogs continue to strongly avoid lions, and consequently avoid habitats associated with their two most important prey species. Although the density of lions in the GKE is low due to prey depletion, their competitive effects on wild dogs remain strong. These effects are likely compounded by prey-base homogenization, as lions in the GKE now rely heavily on the same prey preferred by wild dogs. These results suggest that a reduction in lion density does not necessarily reduce competition, and helps explain why wild dogs decline in parallel with their dominant competitors in ecosystems suffering from anthropogenic prey depletion. Protecting prey populations within the few remaining strongholds for wild dogs is vitally important to avoid substantial population declines. Globally, understanding the impacts of prey depletion on carnivore guild dynamics should be an increasingly important area of focus for conservation.