Ecology

Permanent URI for this communityhttps://scholarworks.montana.edu/handle/1/44

The department's teaching and research addresses critical ecological and natural resources issues for Montana, but also tackles fundamental and applied questions around the globe. Undergraduate programs within the department include Fish & Wildlife Management and Ecology, Conservation Biology and Ecology, Organismal Biology, and Biology Teaching. Graduate programs (M.S. and P.hD.) include Fish & Wildlife Management or Biology and Biological Sciences and an intercollege PhD in Ecology and Environmental Sciences.

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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.
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, Egil
Most 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.
Effects of a protection gradient on carnivore density and survival: an example with leopards in the Luangwa valley, Zambia
(2016-06) Rosenblatt, Elias G.; Creel, Scott; Becker, Matthew S.; Merkle, Johnathan; Mwape, Henry; Schuette, Paul; Simpamba, Twakundine
Human activities on the periphery of protected areas can limit carnivore populations, but measurements of the strength of such effects are limited, largely due to difficulties of obtaining precise data on population density and survival. We measured how density and survival rates of a previously unstudied leopard population varied across a gradient of protection and evaluated which anthropogenic activities accounted for observed patterns. Insights into this generalist's response to human encroachment are likely to identify limiting factors for other sympatric carnivore species. Motion-sensitive cameras were deployed systematically in adjacent, similarly sized, and ecologically similar study areas inside and outside Zambia's South Luangwa National Park (SLNP) from 2012 to 2014. The sites differed primarily in the degree of human impacts: SLNP is strictly protected, but the adjacent area was subject to human encroachment and bushmeat poaching throughout the study, and trophy hunting of leopards prior to 2012. We used photographic capture histories with robust design capture-recapture models to estimate population size and sex-specific survival rates for the two areas. Leopard density within SLNP was 67% greater than in the adjacent area, but annual survival rates and sex ratios did not detectably differ between the sites. Prior research indicated that wire-snare occurrence was 5.2 times greater in the areas adjacent to the park. These results suggest that the low density of leopards on the periphery of SLNP is better explained by prey depletion, rather than by direct anthropogenic mortality. Long-term spatial data from concurrent lion studies suggested that interspecific competition did not produce the observed patterns. Large carnivore populations are often limited by human activities, but science-based management policies depend on methods to rigorously and quantitatively assess threats to populations of concern. Using noninvasive robust design capture-recapture methods, we systematically assessed leopard density and survival across a protection gradient and identified bushmeat poaching as the likely limiting factor. This approach is of broad value to evaluate the impacts of anthropogenic activities on carnivore populations that are distributed across gradients of protection.
Occupancy patterns and niche partitioning within a diverse carnivore community exposed to anthropogenic pressures
(2013-02) Schuette, Paul; Wagner, Aaron P.; Wagner, Meredith E.; Creel, Scott
Although carnivores are in global decline, diverse carnivore communities are common in sub-Saharan Africa, where more than 20 species may co-occur. Though intraguild competition and predation can limit the set of species that coexist, most carnivores have traits that decrease the impacts of interspecific competition on fitness, a pattern that promotes coexistence. An increasing human population and demand for natural resources (e.g. farming) has fragmented landscapes, reduced available prey, and elevated rates of conflict. These anthropogenic pressures tend to eliminate large carnivores first, which can have cascading effects on ecosystem function (e.g. mesopredator release). Anthropogenic pressures might also affect mesocarnivores directly, but this hypothesis has received little research attention to date. Here, we used camera surveys to describe spatial and temporal patterns of carnivore occupancy in a mixed-use landscape in Kenya. This landscape included a community conservation area and seasonally occupied human settlement and livestock grazing areas. We detected 21 carnivore species and examined occupancy patterns for the 12 most frequently detected. Differences among species in responses to environmental conditions supported a hypothesis of spatial niche partitioning. Differences in the temporal activity patterns of the apex predators and mesocarnivores supported a hypothesis of temporal niche partitioning. Human land use altered occupancy patterns in 10 of 12 species. Apex predator occupancies were lower in more anthropogenically disturbed areas, but mesocarnivore occupancies were not inversely related to apex predators, contrary to the mesopredator release hypothesis. Our results suggest that a diverse carnivore community persists in this mixed use landscape because of seasonal variation in human land use.
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.
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.
Carnivore distributions across chaparral habitats exposed to wildfire and rural housing in southern California
(2014-05) Schuette, Paul; Diffendorfer, J. E.; Deutschman, D. H.; Tremor, S.; Spencer, W.
Chaparral and coastal sage scrub habitats in southern California support biologically diverse plant and animal communities. However, native plant and animal species within these shrubland systems are increasingly exposed to human-caused wildfires and an expansion of the human–wildland interface. Few data exist to evaluate the effects of fire and anthropogenic pressures on plant and animal communities found in these environments. This is particularly true for carnivore communities. To address this knowledge gap, we collected detection–non-detection data with motion-sensor cameras and track plots to measure carnivore occupancy patterns following a large, human-caused wildfire (1134 km2) in eastern San Diego County, California, USA, in 2003. Our focal species set included coyote (Canis latrans), gray fox (Urocyon cinereoargenteus), bobcat (Lynx rufus) and striped skunk (Mephitis mephitis). We evaluated the influence on species occupancies of the burned environment (burn edge, burn interior and unburned areas), proximity of rural homes, distance to riparian area and elevation. Gray fox occupancies were the highest overall, followed by striped skunk, coyote and bobcat. The three species considered as habitat and foraging generalists (gray fox, coyote, striped skunk) were common in all conditions. Occupancy patterns were consistent through time for all species except coyote, whose occupancies increased through time. In addition, environmental and anthropogenic variables had weak effects on all four species, and these responses were species-specific. Our results helped to describe a carnivore community exposed to frequent fire and rural human residences, and provide baseline data to inform fire management policy and wildlife management strategies in similar fire-prone ecosystems.
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, Paul
Large 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.
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.

Ecology

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