Publications by Colleges and Departments (MSU - Bozeman)
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Item 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, FredAllocating 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.Item 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, TwakundineFactors 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 (≥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.Item The relationship between direct predation and antipredator responses: a test with multiple predators and multiple prey(2017-08) Creel, Scott; M'soka, Jassiel; Smit, Daan; Becker, Matthew S.; Christianson, David A.; Schuette, Paul; Dröge, EgilMost species adjust their behavior to reduce the likelihood of predation. Many experiments have shown that antipredator responses carry energetic costs that can affect growth, survival, and reproduction, so that the total cost of predation depends on a trade-off between direct predation and risk effects. Despite these patterns, few field studies have examined the relationship between direct predation and the strength of antipredator responses, particularly for complete guilds of predators and prey. We used scan sampling in 344 observation periods over a four-year field study to examine behavioral responses to the immediate presence of predators for a complete antelope guild (dominated by wildebeest, zebra, and oribi) in Liuwa Plains National Park, Zambia, testing for differences in response to all large carnivores in the ecosystem (lions, spotted hyenas, cheetahs, and African wild dogs). We quantified the proportion that each prey species contributed to the kills made by each predator (516 total kills), used distance sampling on systematic line transects to determine the abundance of each prey species, and combined these data to quantify the per-capita risk of direct predation for each predator-prey pair. On average, antelopes increased their vigilance by a factor of 2.4 when predators were present. Vigilance varied strongly among prey species, but weakly in response to different predators. Increased vigilance was correlated with reduced foraging in a similar manner for all prey species. The strength of antipredator response was not detectably related to patterns of direct predation (n = 15 predator-prey combinations with sufficient data). This lack of correlation has implications for our understanding of the role of risk effects as part of the limiting effect of predators on prey.Item Coexistence of African lions, livestock, and people in a landscape with variable human land use and seasonal movements(2013-01) Schuette, Paul; Creel, Scott; Christianson, David A.Apex carnivores around the world have experienced rapid population declines and local extirpation due to anthropogenic pressures, and they are increasingly restricted to government-protected areas (GPAs). Though GPAs are critical for carnivore conservation, mixed-use landscapes may be crucial for sustaining viable populations. Few studies, particularly in Africa, have examined joint use of a landscape by people and conflict-prone carnivores, such as the African lion (Panthera leo), in a situation where conflict is low. In southern Kenya, we studied a lion population in an unfenced rangeland occupied by the Maasai people and their livestock. The Maasai shift their settlements and grazing areas seasonally across a permanent river, a practice we hypothesized might promote coexistence. We radio-collared lions (n = 6) to determine density and document spatial patterns in response to seasonal movements of people in a Conservation Area and buffer zone (250 km2). Despite high livestock density, lion density was comparable to many GPAs (0.136 individuals/km2). Lion spatial distribution and habitat selection shifted in relation to seasonal movements of people and livestock. Conflict was low, likely because lions increased their use of the Conservation Area and dense habitats when people were nearby. Lion responses to human movements reduced access to permanent water, but not prey. A land use system based on temporary settlements and grazing areas allowed lions to co-occur with people and livestock at high density. These results suggest a general strategy for the conservation of apex carnivores outside of GPAs, focusing on areas that exhibit spatiotemporal variation in human land use.Item Conserving large populations of lions - the argument for fences has holes(2013-07-09) Creel, Scott; Becker, Matthew S.; Durant, S. M.; M'soka, Jassiel; Matandiko, Wigganson; Dickman, A. J.; Christianson, David A.; Dröge, E.; Mweetwa, Thandiwe; Pettorelli, N.; Rosenblatt, Elias G.; Schuette, Paul; Woodroffe, R.; Bashir, S.; Beudels-Jamar, R. C.; Blake, S.; Borner, M.; Breitenmoser, C.; Broekhuis, F.; Cozzi, G.; Davenport, T. R. B.; Deutsch, J.; Dollar, L.; Dolrenry, Stephanie; Douglas-Hamilton, I.; Fitzherbert, E.; Foley, C.; Hazzah, L.; Henschel, P.; Hilborn, R.; Hopcraft, J. G. C.; Ikanda, D.; Jacobson, A.; Joubert, B.; Kelly, M. S.; Lichtenfeld, L.; Mace, G. M.; Milanzi, J.; Mitchell, N.; Msuha, M.; Muir, R.; Nyahongo, J.; Pimm, S.; Purchase, G.; Schenck, C.; Sillera-Zubiri, C.; Sinclair, A. R. E.; Songorwa, A. N.; Stanley-Price, M.; Tehou, C. A.; Trout, C.; Wall, J.; Wittemyer, G.; Zimmermann, A.Packer et al. reported that fenced lion populations attain densities closer to carrying capacity than unfenced populations. However, fenced populations are often maintained above carrying capacity, and most are small. Many more lions are conserved per dollar invested in unfenced ecosystems, which avoid the ecological and economic costs of fencing.Item Effects of predation risk on group size, vigilance and foraging behavior in an African ungulate community(2014-08) Creel, Scott; Schutte, Paul; Christianson, David A.Predators alter prey dynamics by direct killing and through the costs of antipredator responses or risk effects. Antipredator behavior includes proactive responses to long-term variation in risk (e.g., grouping patterns) and reactive responses to short-term variation in risk (e.g., intense vigilance). In a 3-year field study, we measured variation in antipredator responses and the foraging costs of these responses for 5 ungulates (zebra, wildebeest, Grant’s gazelle, impala, and giraffe) that comprised more than 90% of the prey community available to the 2 locally dominant predators, lions and spotted hyenas. Using a model-selection approach, we examined how vigilance and group size responded to attributes of the predator, prey, and environment. We found that 1) the strength of antipredator responses was affected by attributes of the predator, prey, and environment in which they met; 2) grouping and vigilance were complementary responses; 3) grouping was a proactive response to the use of dangerous habitats, whereas vigilance was a reactive response to finer cues about predation risk; 4) increased vigilance caused a large reduction in foraging for some species (but not all); and 5) there was no clear relationship between direct predation rates and the foraging costs of antipredator responses. Broadly, our results show that antipredator responses and their costs vary in a complex manner among prey species, the predators they face, and the environment in which they meet.Item Neonatal mortality of elk driven by climate, predator phenology and predator community composition(2011-05) Griffin, K.; Hebblewhite, Mark; Robinson, H.; Zager, Peter; Barber-Meyer, S.; Christianson, David A.; Creel, Scott; Harris, N.; Hurley, M.; Jackson, D.Understanding the interaction among predators and between predation and climate is critical to understanding the mechanisms for compensatory mortality. We used data from 1999 radio-marked neonatal elk (Cervus elaphus) calves from 12 populations in the north-western United States to test for effects of predation on neonatal survival, and whether predation interacted with climate to render mortality compensatory. Weibull survival models with a random effect for each population were fit as a function of the number of predator species in a community (3–5), seven indices of climatic variability, sex, birth date, birth weight, and all interactions between climate and predators. Cumulative incidence functions (CIF) were used to test whether the effects of individual species of predators were additive or compensatory. Neonatal elk survival to 3 months declined following hotter previous summers and increased with higher May precipitation, especially in areas with wolves and/or grizzly bears. Mortality hazards were significantly lower in systems with only coyotes (Canis latrans), cougars (Puma concolor) and black bears (Ursus americanus) compared to higher mortality hazards experienced with gray wolves (Canis lupus) and grizzly bears (Ursus horribilis). In systems with wolves and grizzly bears, mortality by cougars decreased, and predation by bears was the dominant cause of neonatal mortality. Only bear predation appeared additive and occurred earlier than other predators, which may render later mortality by other predators compensatory as calves age. Wolf predation was low and most likely a compensatory source of mortality for neonatal elk calves. Functional redundancy and interspecific competition among predators may combine with the effects of climate on vulnerability to predation to drive compensatory mortality of neonatal elk calves. The exception was the evidence for additive bear predation. These results suggest that effects of predation by recovering wolves on neonatal elk survival, a contentious issue for management of elk populations, may be less important than the composition of the predator community. Future studies would benefit by synthesizing overwinter calf and adult-survival data sets, ideally from experimental studies, to test the roles of predation in annual compensatory and additive mortality of elk.Item A survey of the effects of wolf predation risk on pregnancy rates and calf recruitment in elk(2011-12) Creel, Scott; Christianson, David A.; Winnie, John A. Jr.We have previously found that the behavioral responses of elk to the risk of predation by wolves are correlated with changes in habitat selection, altered diets, reduced food intake, decreased fecal progesterone concentrations, and decreased calf recruitment. P. J. White et al. recently questioned these results, concluding that “multiple lines of evidence and/or strong validation should be brought to bear before relying on indirect measures of how predators affect pregnancy rates.” Here, we systematically surveyed available data and found that five studies (with data from 10 widely distributed populations) have directly detected decreases of 24–43% in elk pregnancy rates in response to increased predation risk. This survey includes data not discussed by White et al. from their own research, which reveal a 32% decrease in pregnancy rates following wolf recolonization in central Yellowstone. Following the survey of available data, we discuss several methodological and statistical problems in White et al.'s study that would be expected to mask the effects of predation risk. While other factors also clearly affect elk recruitment, multiple lines of evidence using a broad array of methods have detected an association between predation risk and reduced reproduction in elk.Item Glucocorticoid stress responses of lions in relation to group composition, human land use and proximity to people(2013-06) Creel, Scott; Christianson, David A.; Schuette, PaulLarge carnivore populations are in global decline, and conflicts between large carnivores and humans or their livestock contribute to low tolerance of large carnivores outside of protected areas. African lions (Panthera leo) are a conflict-prone species, and their continental range has declined by 75% in the face of human pressures. Nonetheless, large carnivore populations persist (or even grow) in some areas that are occupied by humans. Lions attain locally high density in the Olkiramatian and Shompole Group Ranches of Kenya's South Rift region, despite residence by pastoralist Maasai people and their sheep, goats, and cattle. We have previously found that these lions respond to seasonal movements of people by moving away from occupied settlements, shifting into denser habitats when people are nearby, and moving into a protected conservation area when people move into the adjacent buffer zone. Here, we examined lion stress responses to anthropogenic activities, using enzyme-linked immunoassay to measure the concentration of faecal glucocorticoid metabolites in 136 samples collected from five lion groups over 2 years. Faecal glucocorticoid metabolite concentrations were significantly lower for lions in the conservation area than for lions in the human-settled buffer zone, and decreased significantly with increasing distance to the nearest occupied human settlement. Faecal glucocorticoid metabolite concentrations were not detectably related to fine-scaled variation in prey or livestock density, and surprisingly, faecal glucocorticoid metabolite concentrations were higher in the wet season, when regional prey abundance was high. Lions coexist with people and livestock on this landscape by adjusting their movements, but they nonetheless mount an appreciable stress response when conditions do not allow them to maintain adequate separation. Thus, physiological data confirm inferences from prior data on lion movements and habitat use, showing that access to undisturbed and protected areas facilitates human–lion coexistence in a broader landscape that is used by people and livestock.Item Underestimating the frequency, strength and cost of anti-predator responses with data from GPS collars: an example with wolves and elk.(2013-11) Creel, Scott; Winnie, John A. Jr.; Christianson, David A.Field studies that rely on fixes from GPS-collared predators to identify encounters with prey will often underestimate the frequency and strength of antipredator responses. These underestimation biases have several mechanistic causes. (1) Step bias: The distance between successive GPS fixes can be large, and encounters that occur during these intervals go undetected. This bias will generally be strongest for cursorial hunters that can rapidly cover large distances (e.g., wolves and African wild dogs) and when the interval between GPS fixes is long relative to the duration of a hunt. Step bias is amplified as the path travelled between successive GPS fixes deviates from a straight line. (2) Scatter bias: Only a small fraction of the predators in a population typically carry GPS collars, and prey encounters with uncollared predators go undetected unless a collared group-mate is present. This bias will generally be stronger for fission–fusion hunters (e.g., spotted hyenas, wolves, and lions) than for highly cohesive hunters (e.g., African wild dogs), particularly when their group sizes are large. Step bias and scatter bias both cause underestimation of the frequency of antipredator responses. (3) Strength bias: Observations of prey in the absence of GPS fix from a collared predator will generally include a mixture of cases in which predators were truly absent and cases in which predators were present but not detected, which causes underestimation of the strength of antipredator responses. We quantified these biases with data from wolves and African wild dogs and found that fixes from GPS collars at 3-h intervals underestimated the frequency and strength of antipredator responses by a factor >10. We reexamined the results of a recent study of the nonconsumptive effects of wolves on elk in l