Browsing by Author "Paterson, J. Terrill"
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Item Assessing respiratory pathogen communities in bighorn sheep populations: Sampling realities, challenges, and improvements(2017-07) Butler, Carson J.; Edwards, William H.; Jennings-Gaines, Jessica E.; Killion, Halcyon J.; Wood, Mary E.; McWhirter, Douglas E.; Paterson, J. Terrill; Proffitt, Kelly M.; Almberg, Emily S.; White, Patrick J.; Rotella, Jay J.; Garrott, Robert A.Respiratory disease has been a persistent problem for the recovery of bighorn sheep (Ovis canadensis), but has uncertain etiology. The disease has been attributed to several bacterial pathogens including Mycoplasma ovipneumoniae and Pasteurellaceae pathogens belonging to the Mannheimia, Bibersteinia, and Pasteurella genera. We estimated detection probability for these pathogens using protocols with diagnostic tests offered by a fee-for-service laboratory and not offered by a fee-for-service laboratory. We conducted 2861 diagnostic tests on swab samples collected from 476 bighorn sheep captured across Montana and Wyoming to gain inferences regarding detection probability, pathogen prevalence, and the power of different sampling methodologies to detect pathogens in bighorn sheep populations. Estimated detection probability using fee-for-service protocols was less than 0.50 for all Pasteurellaceae and 0.73 for Mycoplasma ovipneumoniae. Non-fee-for-service Pasteurellaceae protocols had higher detection probabilities, but no single protocol increased detection probability of all Pasteurellaceae pathogens to greater than 0.50. At least one protocol resulted in an estimated detection probability of 0.80 for each pathogen except Mannheimia haemolytica, for which the highest detection probability was 0.45. In general, the power to detect Pasteurellaceae pathogens at low prevalence in populations was low unless many animals were sampled or replicate samples were collected per animal. Imperfect detection also resulted in low precision when estimating prevalence for any pathogen. Low and variable detection probabilities for respiratory pathogens using live-sampling protocols may lead to inaccurate conclusions regarding pathogen community dynamics and causes of bighorn sheep respiratory disease epizootics. We recommend that agencies collect multiples samples per animal for Pasteurellaceae detection, and one sample for Mycoplasma ovipneumoniae detection from at least 30 individuals to reliably detect both Pasteurellaceae and Mycoplasma ovipneumoniae at the population-level. Availability of PCR diagnostic tests to wildlife management agencies would improve the ability to reliably detect Pasteurellaceae in bighorn sheep populations.Item Birth dates vary with fixed and dynamic maternal features, offspring sex, and extreme climatic events in a high-latitude marine mammal(2016-02) Rotella, Jay J.; Paterson, J. Terrill; Garrott, Robert A.Reproductive synchrony tends to be widespread in diverse species of plants and animals, especially at higher latitudes. However, for long-lived mammals, birth dates for different individuals can vary by weeks within a population. A mother\'s birth timing can reveal useful information about her reproductive abilities and have important implications for the characteristics and survival of her offspring. Despite this, our current knowledge of factors associated with variation in birth dates is modest. We used long-term data for known-age Weddell seals in Antarctica and a Bayesian hierarchical modeling approach to study how birth dates varied with fixed and temporally varying features of mothers, whether sex allocation varied with birth timing, and annual variation in birth dates. Based on birth dates for 4465 pups born to 1117 mothers aged 4-31, we found that diverse features of mothers were associated with variation in birth dates. Maternal identity was the most important among these. Unlike most studies, which have reported that birth dates occur earlier as mothers age, we found that birth dates progressively occurred earlier in the year in the early part of a mother\'s reproductive life, reached a minimum at age 16, and then occurred later at later ages. Birth dates were positively related to a mother\'s age at primiparity and recent reproductive effort. The earliest birth dates were for pups born to prime-age mothers who did not reproduce in the previous year but began reproduction early in life, suggesting that females in the best condition gave birth earlier than others. If so, our finding that male pups tended to be born earlier than females provides support for the Trivers-Willard sex-allocation model. Average birth dates were quite consistent across years, except for 2 years that had notable delays and occurred during the period when massive icebergs were present and disrupted the ecosystem.Item Drivers of variation in the population dynamics of bighorn sheep(Wiley, 2021-07) Paterson, J. Terrill; Proffitt, Kelly M.; Rotella, Jay; McWhirter, Douglas; Garrott, RobertUnderstanding how variation in vital rates interact to shape the trajectories of populations has long been understood to be a critical component of informed management and restoration efforts. However, an expanding body of work suggests that the expectations for population dynamics of ungulates may not be applicable to small, declining, or threatened populations. Populations of bighorn sheep (Ovis canadensis) suffered declines at the turn of the 20th century, and restoration efforts have been mixed such that many populations remain small and isolated. Here, we utilized survey data collected from 1983 to 2018 from 17 populations of bighorn sheep in Montana and Wyoming to estimate the parameters of a stage-specific population model that we used to (1) characterize the spatial and temporal variation in key vital rates including whether populations were stable, increasing, or declining; (2) estimate the contributions of vital rates to variation in population growth rates; and (3) evaluate potential sources of variation in lamb survival. We found substantial variation in all vital rates both among years and populations, strong evidence for an overall decline in nine of the 17 populations, and clear evidence for multiple combinations of vital rates that resulted in positive population trajectories. The contribution of ewe survival and lamb survival to the total variation in population growth rates varied among populations; however, declines in ewe survival dominated transitions of population trajectories from stable or increasing to declining, whereas reversals of declining population trajectories were dominated by improved lamb survival. We found strong evidence for a diverse set of associations between lamb survival and environmental covariates related to growing season and winter severity. The estimated relationships predict that environmental drivers can cause important changes in lamb survival and provide suggestive evidence that the presence of Mycoplasma ovipneumoniae is not sufficient to prevent population growth. Although our work demonstrates that the trajectories of these populations of bighorn sheep are driven by a variety of processes, the diversity of relationships between vital rates and population growth rates also suggests that there are multiple pathways to manage for population recovery.Item Evaluating the importance of individual heterogeneity in reproduction to Weddell seal population dynamics using integral projection models(Wiley, 2023-07) Macdonald, Kaitlin R.; Rotella, Jay J.; Paterson, J. TerrillIdentifying and accounting for unobserved individual heterogeneity in vital rates in demographic models is important for estimating population-level vital rates and identifying diverse life-history strategies, but much less is known about how this individual heterogeneity influences population dynamics. We aimed to understand how the distribution of individual heterogeneity in reproductive and survival rates influenced population dynamics using vital rates from a Weddell seal population by altering the distribution of individual heterogeneity in reproduction, which also altered the distribution of individual survival rates through the incorporation of our estimate of the correlation between the two rates and assessing resulting changes in population growth. We constructed an integral projection model (IPM) structured by age and reproductive state using estimates of vital rates for a long-lived mammal that has recently been shown to exhibit large individual heterogeneity in reproduction. Using output from the IPM, we evaluated how population dynamics changed with different underlying distributions of unobserved individual heterogeneity in reproduction. Results indicate that the changes to the underlying distribution of individual heterogeneity in reproduction cause very small changes in the population growth rate and other population metrics. The largest difference in the estimated population growth rate resulting from changes to the underlying distribution of individual heterogeneity was less than 1%. Our work highlights the differing importance of individual heterogeneity at the population level compared to the individual level. Although individual heterogeneity in reproduction may result in large differences in the lifetime fitness of individuals, changing the proportion of above- or below-average breeders in the population results in much smaller differences in annual population growth rate. For a long-lived mammal with stable and high adult-survival that gives birth to a single offspring, individual heterogeneity in reproduction has a limited effect on population dynamics. We posit that the limited effect of individual heterogeneity on population dynamics may be due to canalization of life-history traits.Item Evaluating the summer landscapes of predation risk and forage quality for elk ( Cervus canadensis )(Wiley, 2022-08) Paterson, J. Terrill; Proffitt, Kelly M.; DeCesare, Nicholas J.; Gude, Justin A.; Hebblewhite, MarkThe recovery of carnivore populations in North American has consequences for trophic interactions and population dynamics of prey. In addition to direct effects on prey populations through killing, predators can influence prey behavior by imposing the risk of predation. The mechanisms through which patterns of space use by predators are linked to behavioral response by prey and nonconsumptive effects on prey population dynamics are poorly understood. Our goal was to characterize population- and individual-level patterns of resource selection by elk (Cervus canadensis) in response to risk of wolves (Canis lupus) and mountain lions (Puma concolor) and evaluate potential nonconsumptive effects of these behavioral patterns. We tested the hypothesis that individual elk risk-avoidance behavior during summer would result in exposure to lower-quality forage and reduced body fat and pregnancy rates. First, we evaluated individuals' second-order and third-order resource selection with a used-available sampling design. At the population level, we found evidence for a positive relationship between second- and third-order selection and forage, and an interaction between forage quality and mountain lion risk such that the relative probability of use at low mountain lion risk increased with forage quality but decreased at high risk at both orders of selection. We found no evidence of a population-level trade-off between forage quality and wolf risk. However, we found substantial among-individual heterogeneity in resource selection patterns such that population-level patterns were potentially misleading. We found no evidence that the diversity of individual resource selection patterns varied predictably with available resources, or that patterns of individual risk-related resource selection translated into biologically meaningful changes in body fat or pregnancy rates. Our work highlights the importance of evaluating individual responses to predation risk and predator hunting technique when assessing responses to predators and suggests nonconsumptive effects are not operating at a population scale in this system.Item Investigating diverse sources of variation in the amount of time Weddell seal ( Leptonychotes weddellii ) pups spend in the water during the lactation period(Wiley, 2022-07) Petch, Shane M.; Rotella, Jay J.; Link, William A.; Paterson, J. Terrill; Garrott, Robert A.Field observations suggest that time spent in the water by Weddell seal pups during lactation varies among individuals, which could yield important developmental tradeoffs. We analyzed data from 713 pups born to 419 different mothers over 9 years to evaluate total time in the water, age at first entry, and potential sources of variation using temperature loggers attached to the rear flipper of pups. Pups first entered the water at 11–29 days of age (M = 14.9) and spent 4–204 hr (M = 69.3) in the water by 30 days of age. Age at first entry was earlier for pups with higher birth mass and mothers of above average reproductive experience. Total time in the water was related to maternal identity and greater for female pups and for pups that had higher birth mass, mothers of intermediate age, mothers that skipped reproduction in the previous year, and for pups that first entered the water at younger ages. Phenotypic traits explain observed variation in the development of a key life history behavior in the Weddell seal. Strong individual variation in time spent in metabolically costly swimming and diving might lead to variation in growth, energy stores, and survival and fitness outcomes.Item Patterns of age-related change in reproductive effort differ in the pre-natal and post-natal periods in a long-lived mammal.(2016-09) Paterson, J. Terrill; Rotella, Jay J.; Mannas, Jennifer M.; Garrott, Robert A.Age-related changes in maternal reproductive allocation for long-lived species are a key prediction from life-history theory. Theoretical and empirical work suggests that allocation may increase with age due to constraint (increases with experience) or restraint (increases with age in the face of declining residual reproductive value), and may decrease among the oldest aged animals due to senescence in reproductive function. Here, we use a hierarchical modelling approach to investigate the age-related patterns of change in maternal reproductive effort in the Weddell seal, a long-lived marine mammal with a protracted period of maternal care during which mothers allocate a large proportion of body mass while feeding little. We find that maternal allocation increases with age for young mothers during both the pre-natal and post-natal periods. In contrast, older mothers demonstrate a senescent decline in pre-natal allocation but allocate more of their declining resources to their offspring during the post-natal period. We also find strong evidence for the importance of individual effects in reproductive allocation among mothers: some mothers consistently produce heavier (or lighter) pups than expected. Our results indicate that maternal allocation changes over a mother\'s reproductive life span and that age-specific differences differ in notable ways in pre-natal and post-natal periods.Item Respiratory pathogens and their association with population performance in Montana and Wyoming bighorn sheep populations(2018-11) Butler, Carson J.; Edwards, William H.; Paterson, J. Terrill; Proffitt, Kelly M.; Jennings-Gaines, Jessica E.; Killion, Halcyon J.; Wood, Mary E.; Ramsey, Jennifer M.; Almberg, Emily S.; Dewey, Sarah R.; McWhirter, Douglas E.; Courtemanch, Alyson B.; White, Patrick J.; Rotella, Jay J.; Garrott, Robert A.At the request of National Park Service resource managers, we began a study in 2000 to evaluate causes for the decline of the bighorn sheep (Ovis canadensis) population inhabiting Bighorn Canyon National Recreation Area (BICA), the Pryor Mountain Wild Horse Range, and surrounding state and U.S. Forest Service lands in Montana and Wyoming. Our study consisted of radio-collaring adult rams and ewes with mortality sensors to monitor adult mortalities, tracking ewes to determine pregnancy and lambing rates, habitat assessments to determine why the population was not expanding into what had been modeled using GIS methodology as suitable bighorn sheep habitat, measuring ungulate herbaceous consumption rates and herbaceous production to determine plant responses, and aerial and boat surveys to determine bighorn sheep population range and population dynamics (Schoenecker and others, this report). Two habitat suitability models were created and conducted (Gudorf, this report; Wockner and others, this report) using different methodologies, and comparisons made between the two. Herd population dynamics were modeled using the POP-II and POP-III programs (Roelle, this report), and a reassessment of ungulate exclosures that were established 8–10 years ago was conducted (Gerhardt, this report). The bighorn sheep population of the greater Bighorn Canyon National Recreation Area (BICA) was extirpated in the 1800s, and then reintroduced in 1973. The herd increased to a peak population of about 211 animals (Kissell and others, 1996), but then declined sharply in 1995 and 1996. Causes for the decline were unknown. Numbers have remained around 100 ± 20 animals since 1998. Previous modeling efforts determined what areas were suitable bighorn sheep habitat (Gudorf and others, 1996). We tried to determine why sheep were not using areas that were modeled as suitable or acceptable habitat, and to evaluate population dynamics of the herd.Item Simulation-based validation of spatial capture-recapture models: A case study using mountain lions(2019-04) Paterson, J. Terrill; Proffitt, Kelly M.; Jimenez, Ben; Rotella, Jay J.; Garrott, Robert A.Spatial capture-recapture (SCR) models have improved the ability to estimate densities of rare and elusive animals. However, SCR models have seldom been validated even as model formulations diversify and expand to incorporate new sampling methods and/or additional sources of information on model parameters. Information on the relationship between encounter probabilities, sources of additional information, and the reliability of density estimates, is rare but crucial to assessing reliability of SCR-based estimates. We used a simulation-based approach that incorporated prior empirical work to assess the accuracy and precision of density estimates from SCR models using spatially unstructured sampling. To assess the consequences of sparse data and potential sources of bias, we simulated data under six scenarios corresponding to three different levels of search effort and two levels of correlation between search effort and animal density. We then estimated density for each scenario using four models that included increasing amounts of information from harvested individuals and telemetry to evaluate the impact of additional sources of information. Model results were sensitive to the quantity of available information: density estimates based on low search effort were biased high and imprecise, whereas estimates based on high search effort were unbiased and precise. A correlation between search effort and animal density resulted in a positive bias in density estimates, though the bias decreased with increasingly informative datasets. Adding information from harvested individuals and telemetered individuals improved density estimates based on low and moderate effort but had negligible impact for datasets resulting from high effort. We demonstrated that density estimates from SCR models using spatially unstructured sampling are reliable when sufficient information is provided. Accurate density estimates can result if empirical-based simulations such as those presented here are used to develop study designs with appropriate amounts of effort and information sources.Item Tight coupling of primary production and marine mammal reproduction in the Southern Ocean(2015-05) Paterson, J. Terrill; Rotella, Jay J.; Arrigo, Kevin; Garrott, Robert A.Polynyas are areas of open water surrounded by sea ice and are important sources of primary production in high-latitude marine ecosystems. The magnitude of annual primary production in polynyas is controlled by the amount of exposure to solar radiation and sensitivity to changes in sea-ice extent. The degree of coupling between primary production and production by upper trophic-level consumers in these environments is not well understood, which prevents reliable predictions about population trajectories for species at higher trophic levels under potential future climate scenarios. In this study, we find a strong, positive relationship between annual primary production in an Antarctic polynya and pup production by ice-dependent Weddell seals. The timing of the relationship suggests reproductive effort increases to take advantage of high primary production occurring in the months after the birth pulse. Though the proximate causal mechanism is unknown, our results indicate tight coupling between organisms at disparate trophic levels on a short timescale, deepen our understanding of marine ecosystem processes, and raise interesting questions about why such coupling exists and what implications it has for understanding high-latitude ecosystems.Item Variation in the vital rates of an Antarctic marine predator: the role of individual heterogeneity(2018-10) Paterson, J. Terrill; Rotella, Jay J.; Link, William A.; Garrott, Robert A.Variation in life-history traits such as lifespan and lifetime reproductive output is thought to arise, in part, due to among-individual differences in the underlying probabilities of survival and reproduction. However, the stochastic nature of demographic processes can also generate considerable variation in fitness-related traits among otherwise-identical individuals. An improved understanding of life-history evolution and population dynamics therefore depends on evaluating the relative role of each of these processes. Here, we used a 33-yr data set with reproductive histories for 1,274 female Weddell seals from Erebus Bay, Antarctica, to assess the strength of evidence for among-individual heterogeneity in the probabilities of survival and reproduction, while accounting for multiple other sources of variation in vital rates. Our analysis used recent advances in Bayesian model selection techniques and diagnostics to directly compare model fit and predictive power between models that included individual effects on survival and reproduction to those that did not. We found strong evidence for costs of reproduction to both survival and future reproduction, with breeders having rates of survival and subsequent reproduction that were 3% and 6% lower than rates for non-breeders. We detected age-related changes in the rates of survival and reproduction, but the patterns differed for the two rates. Survival rates steadily declined from 0.92 at age 7 to 0.56 at the maximal age of 31yr. In contrast, reproductive rates increased from 0.68 at age 7 to 0.79 at age 16 and then steadily declined to 0.37 for the oldest females. Models that included individual effects explained more variation in observed life histories and had better estimated predictive power than those that did not, indicating their importance in understanding sources of variation among individuals in life-history traits. We found that among-individual heterogeneity in survival was small relative to that for reproduction. Our study, which found patterns of variation in vital rates that are consistent with a series of predictionsfrom life-history theory, is the first to provide a thorough assessment of variation in important vital rates for a long-lived, high-latitude marine mammal while taking full advantage of recent developments in model evaluation.