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| 2. |
- Creel, Scott
(författare)
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A retrospective view of early research on dominance, stress and reproduction in cooperatively breeding carnivores
- 2022
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Ingår i: Hormones and Behavior. - : Elsevier BV. - 0018-506X .- 1095-6867. ; 140
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Tidskriftsartikel (refereegranskat)abstract
- Social carnivores have been central in studies of cooperative breeding, and research using noninvasive methods to examine behavioral and endocrine mechanisms of reproductive suppression started in the 1980s with dwarf mongooses in Serengeti National Park. Here, I synthesize the methods, findings and limitations of a research program that examined relationships between social dominance, age, mass, aggression, mating, gonadal steroids, glucocorticoids and reproduction in female and male dwarf mongooses, African wild dogs and wolves. Infanticide is a reliable backstop for reproductive suppression in females, and reproduction is energetically costly in these species. These conditions favor hypothalamic - pituitary - gonadal (HPG) adaptations that reduce the fertility of subordinate females to avoid the cost of producing doomed offspring. Infanticide also favors close synchronization of reproduction when subordinate females do become pregnant. In males, infanticide is a less reliable backstop and reproduction is less costly, so direct effects of subordination on fertility are less pronounced. Age is a strong predictor of social dominance in these species, but the evolutionary reason for this is not clear. In dwarf mongooses and wild dogs, alpha females were never deposed by younger packmates, but alpha males were: this difference is also not understood. Patterns of reproduction supported models predicting that alphas are less likely to share reproduction when the fitness costs of reproduction are high, when the fitness expected for dispersers is low, and with young subordinates to whom they are more closely related. Correlations between dominance and adrenal glucocorticoid concentrations varied between species and sexes, but did not support the hypothesis that chronic stress causes reproductive suppression.
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| 3. |
- Creel, Scott
(författare)
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African wild dog movements show contrasting responses to long and short term risk of encountering lions: analysis using dynamic Brownian bridge movement models
- 2022
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Ingår i: Movement Ecology. - : Springer Science and Business Media LLC. - 2051-3933. ; 10
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Tidskriftsartikel (refereegranskat)abstract
- Background Prey depletion is a threat to the world's large carnivores, and is likely to affect subordinate competitors within the large carnivore guild disproportionately. African lions limit African wild dog populations through interference competition and intraguild predation. When lion density is reduced as a result of prey depletion, wild dogs are not competitively released, and their population density remains low. Research examining distributions has demonstrated spatial avoidance of lions by wild dogs, but the effects of lions on patterns of movement have not been tested. Movement is one of the most energetically costly activities for many species and is particularly costly for cursorial hunters like wild dogs. Therefore, testing how top-down, bottom-up, and anthropogenic variables affect movement patterns can provide insight into mechanisms that limit wild dogs (and other subordinate competitors) in resource-depleted ecosystems. Methods We measured movement rates using the motion variance from dynamic Brownian Bridge Movement Models (dBBMMs) fit to data from GPS-collared wild dogs, then used a generalized linear model to test for effects on movement of predation risk from lions, predictors of prey density, and anthropogenic and seasonal variables. Results Wild dogs proactively reduced movement in areas with high lion density, but reactively increased movement when lions were immediately nearby. Predictors of prey density had consistently weaker effects on movement than lions did, but movements were reduced in the wet season and when dependent offspring were present. Conclusion Wild dogs alter their patterns of movement in response to lions in ways that are likely to have important energetic consequences. Our results support the recent suggestion that competitive limitation of wild dogs by lions remains strong in ecosystems where lion and wild dog densities are both low as a result of anthropogenic prey depletion. Our results reinforce an emerging pattern that movements often show contrasting responses to long-term and short-term variation in predation risk.
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| 4. |
- Creel, Scott
(författare)
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Assessing the performance of index calibration survey methods to monitor populations of wide-ranging low-density carnivores
- 2020
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Ingår i: Ecology and Evolution. - : Wiley. - 2045-7758. ; 10, s. 3276-3292
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Tidskriftsartikel (refereegranskat)abstract
- 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.
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| 5. |
- Creel, Scott, et al.
(författare)
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Carnivores, competition and genetic connectivity in the Anthropocene
- 2019
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Ingår i: Scientific Reports. - : Nature Publishing Group. - 2045-2322. ; 9
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Tidskriftsartikel (refereegranskat)abstract
- Current extinction rates are comparable to five prior mass extinctions in the earth's history, and are strongly affected by human activities that have modified more than half of the earth's terrestrial surface. Increasing human activity restricts animal movements and isolates formerly connected populations, a particular concern for the conservation of large carnivores, but no prior research has used high throughput sequencing in a standardized manner to examine genetic connectivity for multiple species of large carnivores and multiple ecosystems. Here, we used RAD SNP genotypes to test for differences in connectivity between multiple ecosystems for African wild dogs (Lycaon pictus) and lions (Panthera leo), and to test correlations between genetic distance, geographic distance and landscape resistance due to human activity. We found weaker connectivity, a stronger correlation between genetic distance and geographic distance, and a stronger correlation between genetic distance and landscape resistance for lions than for wild dogs, and propose a new hypothesis that adaptations to interspecific competition may help to explain differences in vulnerability to isolation by humans.
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| 6. |
- Creel, Scott
(författare)
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Changes in African large carnivore diets over the past half-century reveal the loss of large prey
- 2018
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Ingår i: Journal of Applied Ecology. - : Wiley. - 0021-8901 .- 1365-2664. ; 55, s. 2908-2916
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Tidskriftsartikel (refereegranskat)abstract
- 1. Globally, large carnivores are declining due to direct persecution, habitat loss, and prey depletion. The effects of prey depletion could be amplified by changes in the composition of the herbivore (prey) community that provoke changes in carnivore diets, but this possibility has received little attention.2. We tested for changes over the past half-century in prey selection by the large carnivore guild in Zambia's Kafue National Park (KNP).3. Across 52 predator-prey dyads, 71% of the observed changes showed that large prey have become less important and small prey have become more important. Consequently, dietary niche breadth has decreased for KNP carnivores and niche overlap has increased.4. We tested whether changes in the importance of prey species are related to their current abundance and uniformly found that prey that have increased in importance are now relatively common, while those that have decreased in importance are now relatively rare. We identify four potential effects of these changes for conservation (through intraguild competition, group size, the energetics of hunting, and vulnerability to snaring) that warrant investigation.5. Synthesis and applications. Patterns of prey selection by the large carnivores in Kafue National Park (KNP) have changed appreciably over the past half-century. Decreased predation on large prey, which are now relatively rare, has caused niche compression and increased overlap in carnivore diets. Predation by all KNP large carnivores now concentrates on four small prey species that remain relatively abundant (impala, puku, lechwe, and warthog). Methods to detect such changes in interactions between species are well-established, but are rarely applied to large carnivore-ungulate systems. To guide conservation of ecosystem function, monitoring programmes should consider whether prey depletion alters the patterns of predation or competition within the predator guild because these interactions strongly affect the distribution and abundance of both predators and prey. If the patterns seen in KNP are general, then where carnivores are limited by prey depletion, conservation efforts will be most effective if they focus on mitigating the loss of large prey. In KNP, targeted efforts to protect prey larger than 200 kg, particularly buffalo, should be a priority.
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| 7. |
- Creel, Scott
(författare)
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Hidden Markov Models reveal a clear human footprint on the movements of highly mobile African wild dogs
- 2020
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Ingår i: Scientific Reports. - : Springer Science and Business Media LLC. - 2045-2322. ; 10
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Tidskriftsartikel (refereegranskat)abstract
- Large carnivores have experienced considerable range contraction, increasing the importance of movement across human-altered landscapes between small, isolated populations. African wild dogs (Lycaon pictus) are exceptionally wide-ranging, and recolonization is an important element of their persistence at broad scales. The competition-movement-connection hypothesis suggests that adaptations to move through areas that are unfavorable due to dominant competitors might promote the ability of subordinate competitors (like wild dogs) to move through areas that are unfavorable due to humans. Here, we used hidden Markov models to test how wild dog movements were affected by the Human Footprint Index in areas inside and outside of South Luangwa National Park. Movements were faster and more directed when outside the National Park, but slowed where the human footprint was stronger. Our results can be directly and quantitatively applied to connectivity planning, and we use them to identify ways to better understand differences between species in recent loss of connectivity.
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| 8. |
- Creel, Scott
(författare)
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Hot or hungry? A tipping point in the effect of prey depletion on African wild dogs
- 2023
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Ingår i: Biological Conservation. - 0006-3207 .- 1873-2917. ; 282
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Tidskriftsartikel (refereegranskat)abstract
- Interspecific competition has strong effects within carnivore guilds, and African wild dogs are strongly limited by intraguild predation by lions and food loss to spotted hyenas. The densities of these dominant competitors correlate tightly with prey density, and prey depletion due to snaring is contributing to declines of apex carnivores across Africa. As a consequence of snaring, subordinate competitors like the African wild dog and cheetah are simultaneously experiencing a costly reduction in prey and a beneficial reduction of competitors. The net effect is difficult to predict. Here, we describe a previously unrecognized tipping point in the effect of prey density on wild dog density. Above this tipping point, wild dog density increases as prey density decreases, because the benefits of competitive release outweigh the costs of prey depletion. Below this tipping point, wild dog density decreases as prey density decreases, because the costs of prey depletion outweigh the benefits of competitive release. Conservation strategies for wild dogs must recognize this tipping point. As prey depletion due to snaring becomes severe, it limits wild dogs (mesocarnivores), just as it does lions and spotted hyenas (apex carnivores). Recent analyses based on time trends have argued that climate change is likely to cause wild dog populations to collapse. We suggest that prey depletion provides a mechanistically direct explanation of patterns in wild dog dynamics that have been attributed to climate change, and that the effects of prey depletion are likely to go unrecognized in the absence of data on trends in prey density.
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| 9. |
- Creel, Scott, et al.
(författare)
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Leopard Panthera pardus density and survival in an ecosystem with depressed abundance of prey and dominant competitors
- 2022
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Ingår i: Oryx. - 0030-6053 .- 1365-3008. ; 56, s. 518-527
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Tidskriftsartikel (refereegranskat)abstract
- The leopard Panthera pardus is in range-wide decline, and many populations are highly threatened. Prey depletion is a major cause of global carnivore declines, but the response of leopard survival and density to this threat is unclear: by reducing the density of a dominant competitor (the lion Panthera leo) prey depletion could create both costs and benefits for subordinate competitors. We used capture-recapture models fitted to data from a 7-year camera-trap study in Kafue National Park, Zambia, to obtain baseline estimates of leopard population density and sex-specific apparent survival rates. Kafue is affected by prey depletion, and densities of large herbivores preferred by lions have declined more than the densities of smaller herbivores preferred by leopards. Lion density is consequently low. Estimates of leopard density were comparable to ecosystems with more intensive protection and favourable prey densities. However, our study site is located in an area with good ecological conditions and high levels of protection relative to other portions of the ecosystem, so extrapolating our estimates across the Park or into adjacent Game Management Areas would not be valid. Our results show that leopard density and survival within north-central Kafue remain good despite prey depletion, perhaps because (1) prey depletion has had weaker effects on preferred leopard prey compared to larger prey preferred by lions, and (2) the density of dominant competitors is consequently low. Our results show that the effects of prey depletion can be more complex than uniform decline of all large carnivore species, and warrant further investigation.
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| 10. |
- Creel, Scott
(författare)
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Long-term data reveal fitness costs of anthropogenic prey depletion for a subordinate competitor, the African wild dog (Lycaon pictus)
- 2024
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Ingår i: Ecology and Evolution. - 2045-7758. ; 14
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Tidskriftsartikel (refereegranskat)abstract
- Within carnivore guilds, dominant competitors (e.g., lions, Panthera leo) are limited primarily by the density of prey, while subordinate competitors (e.g., African wild dogs, Lycaon pictus) have been limited by the density of dominant competitors. Historically, the fitness and population density of subordinate competitors have not been tightly linked to prey density. However, populations of large herbivores have declined substantially across sub-Saharan Africa due to human impacts, and where prey depletion is severe, fitness costs for competitive subordinates may begin to outweigh the benefits of competitive release. Using long-term intensive monitoring of African wild dogs in Zambia's Luangwa Valley Ecosystem (LVE), we tested the effects of prey depletion on survival and reproduction. We hypothesized that African wild dog fitness would be lower in prey-depleted areas, despite lower lion densities. Our study area included four contiguous regions that varied in protection level, prey density, and lion density. We fit Bayesian Cormack-Jolly-Seber and closed-capture models to estimate effects on survival and population density, and generalized linear models to estimate effects on reproductive success. We found that the LVE is a stronghold for African wild dogs, with an estimated median density of 4.0 individuals/100 km(2). Despite this high density, survival and reproduction differed among regions, and both components of fitness were substantially reduced in the region with the lowest prey density. Anthropogenic prey depletion is becoming an important limiting factor for African wild dogs. If prey depletion (or any other form of habitat degradation) becomes severe enough that its fitness costs outweigh the benefits of competitive release, such changes can fundamentally alter the balance between limiting factors for competitively subordinate species.
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