College of Letters & Science
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The College of Letters and Science, the largest center for learning, teaching and research at Montana State University, offers students an excellent liberal arts and sciences education in nearly 50 majors, 25 minors and over 25 graduate degrees within the four areas of the humanities, natural sciences, mathematics and social sciences.
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Item 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.Item Effects of de-snaring on the demography and population dynamics of African lions(Elsevier BV, 2023-10) Banda, Kambwiri; Creel, Scott; Sichande, Mwamba; Mweetwa, Thandiwe; Mwape, Henry; Reyes de Merkle, Johnathan; Mwansa Bwalya, Lengwe; Simpamba, Twakundine; McRobb, Rachel; Becker, Matthew S.Lions and other African large carnivores are in decline, due in part to effects of illegal hunting with snares, which can reduce prey availability and directly kill or injure carnivores. It is difficult to effectively remove snares from large ecosystems by patrolling, but an additional approach to reduce effects on large carnivores is to monitor the population closely and de-snare individuals who are found in a snare or have broken free but still carry the wire (often with serious injury). The effectiveness of de-snaring programs to reduce impacts on large carnivores has not been directly tested. Here, we used long-term demographic data from 386 individually identified lions in the Luangwa Valley Ecosystem to test the effects on population growth (λ) and population size (N) of a program to remove snares from injured lions and treat their wounds. Stochastic Leslie matrix projections for a period of five years showed that the population grew with the benefits of de-snaring, but was expected to decline without de-snaring. Mean annual growth (λ) with de-snaring was 1.037 (with growth in 70% of years), closely matching observed changes in population size. Mean annual growth was 0.99 (with growth in 47% of years) for a model that assumed snared animals would have died if not treated, and 0.95 (with growth in 37% of years) for models that also accounted for super-additive effects via the death of dependent cubs and increased infanticide with increased male mortality. De-snaring requires intensive effort, but it can appreciably reduce the effect of snaring on lion population dynamics.Item 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, BridgetLarge 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.Item African wild dog movements show contrasting responses to long and short term risk of encountering lions: analysis using dynamic Brownian bridge movement models(Springer Science and Business Media LLC, 2022-12) Goodheart, Ben; Creel, Scott; Vinks, Milan A.; Banda, Kambwiri; Reyes de Merkle, Johnathan; Kusler, Anna; Dart, Chase; Banda, Kachama; Becker, Matthew S.; Indala, Peter; Simukonda, Chuma; Kaluka, AdrianBackground 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.Item A retrospective view of early research on dominance, stress and reproduction in cooperatively breeding carnivores(Elsevier BV, 2022-04) Creel, ScottSocial 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.Item A retrospective view of early research on dominance, stress and reproduction in cooperatively breeding carnivores(Elsevier BV, 2022-04) Creel, ScottSocial 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. Infanti-cide 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 pro-nounced. 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.Item A retrospective view of early research on dominance, stress and reproduction in cooperatively breeding carnivores(Elsevier BV, 2022-03) Creel, ScottSocial 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.Item 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.