John Winnie - Academia.edu (original) (raw)

Papers by John Winnie

Wildlife Biology

Density is a fundamental ecological parameter that is often difficult to estimate in large herbiv... more Density is a fundamental ecological parameter that is often difficult to estimate in large herbivores due to the wide spatial extent of their populations and imperfect detectability of individuals. We searched Google Earth imagery using an independent double‐observer protocol to estimate elk Cervus elaphus density in Yellowstone National Park, WY, USA, on summer range that had never been systematically surveyed. Detectability of elk was high at 0.836 (95% CRI: 0.769, 0.909) producing high precision in our density estimates. Elk density estimates ranged from a minimum of 8.9 elk km−2 over a 214.8 km2 search area (95% CRI: 7.9, 10.9) to a maximum of 14.0 elk km−2 over 98.5 km2 of nonforested, alpine terrain (95% CRI: 13.5, 14.9). These estimates of elk density are comparable to the highest elk densities ever recorded in Yellowstone which primarily derive from winter, at low elevations, prior to wolf Canis lupus reintroduction. A log‐linear positive effect of open landcover on density ...

These vegetation and fecal data were used to describe the relationship between fecal and vegetati... more These vegetation and fecal data were used to describe the relationship between fecal and vegetation Nitrogen (N) and Phosphorous (P) content, in 2002-2003 in Kruger National Park, as described in Ryan et al, 201

Many studies of mammalian herbivores have employed remotely sensed vegetation greenness, in the f... more Many studies of mammalian herbivores have employed remotely sensed vegetation greenness, in the form of Normalized Difference Vegetation Index (NDVI) as a proxy for forage quality. The assumption that reflected greenness represents forage quality often goes untested, and limited data exist on the relationships between remotely sensed and traditional forage nutrient indicators. We provide the first study connecting NDVI and forage nutrient indicators within a free-ranging African herbivore ecosystem. We examined the relationships between fecal nutrient levels (nitrogen and phosphorus), forage nutrient levels, body condition, and NDVI for African buffalo (Syncerus caffer) in a South African savanna ecosystem over a 2-year period (2001 and 2002). We used an information-theoretic approach to rank models of fecal nitrogen (Nf) and phosphorus (Pf) as functions of geology, season, and NDVI in each year separately. For each year, the highest ranked models for Nf accounted for 61% and 65% of the observed variance, and these models included geology, season, and NDVI. The top-ranked model for Pf in 2001, although capturing 54% of the variability, did not include NDVI. In 2002, we could not identify a top ranking model for phosphorus (i.e., all models were within 2 AICc of each other). Body condition was most highly correlated (equation image; P ≤ 0.001) with NDVI at a 1 month time lag and with Nf at a 3 months time lag (equation image; P ≤ 0.001), but was not significantly correlated with Pf. Our findings suggest that NDVI can be used to index nitrogen content of forage and is correlated with improved body condition in African buffalo. Thus, NDVI provides a useful means to assess forage quality where crude protein is a limiting resource. We found that NDVI accounted for more than a seasonal effect, and in a system where standing biomass may be high but of low quality, understanding available nutrients is useful for management

Ecology and evolution, 2013

Field studies that rely on fixes from GPS-collared predators to identify encounters with prey wil... more 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 ...

Science, 2007

Elk (Cervus elaphus) in the Greater Yellowstone Ecosystem alter patterns of aggregation, habitat ... more Elk (Cervus elaphus) in the Greater Yellowstone Ecosystem alter patterns of aggregation, habitat selection, vigilance, and foraging in the presence of wolves (Canis lupus). Antipredator behaviors like these can reduce predation risk but are also likely to carry costs. Data from five elk populations studied for 16 site years showed that progesterone concentrations (from 1489 fecal samples) declined with the ratio of elk to wolves. In turn, progesterone concentrations were a good predictor of calf recruitment in the subsequent year. Together, these data suggest that wolves indirectly affect the reproductive physiology and the demography of elk through the costs of antipredator behavior.

The Journal of Wildlife Management, 2012

The Journal of Wildlife Management, 2010

Estimating population size in a mark-recapture framework using DNA obtained from remotely collect... more Estimating population size in a mark-recapture framework using DNA obtained from remotely collected genetic samples (e.g., feces) has become common in recent years but rarely has been used for ungulates. Using DNA extracted from fecal pellets, we estimated the size of an argali (Ovis ammon) population that was believed to be isolated from others within the Big Pamir Mountains, Afghanistan, an area where access was difficult and expensive. We used closed-capture models to estimate abundance, and Pradel models to examine closure assumptions, both as implemented in Program MARK. We also made visual counts of argali in the Big Pamirs, allowing comparison of count indices of abundance with modeled estimates. Our model-averaged estimate for female argali in the Big Pamir was 172 (95% CI 5 117-232), which was about 23% higher than our best assessment using uncorrected visual counts. However, mark-recapture models suggested that males were not a closed population; thus, we were unable to provide a meaningful estimate of overall population size. Males either suffered much higher mortality than females during the sampling period, or, more likely, males moved in and out of the Big Pamir area. Although information from DNA did not provide a clear overall population estimate, it suggested that the Big Pamir was not isolated from other argali populations, which could not have been confirmed with visual observations alone. Estimating argali population size using mark-recapture models and fecal DNA is feasible but may be too expensive for frequent monitoring of large and remote populations. Our study demonstrates the importance of sex identification and separate abundance estimation for each sex, especially if movement ecology differs by sex.

Ecology, 2008

Top‐down effects of predators on prey behavior and population dynamics have been extensively stud... more Top‐down effects of predators on prey behavior and population dynamics have been extensively studied. However, some populations of very large herbivores appear to be regulated primarily from the bottom up. Given the importance of food resources to these large herbivores, it is reasonable to expect that forage heterogeneity (variation in quality and quantity) affects individual and group behaviors as well as distribution on the landscape. Forage heterogeneity is often strongly driven by underlying soils, so substrate characteristics may indirectly drive herbivore behavior and distribution. Forage heterogeneity may further interact with predation risk to influence prey behavior and distribution. Here we examine differences in spatial distribution, home range size, and grouping behaviors of African buffalo as they relate to geologic substrate (granite and basalt) and variation in food quality and quantity. In this study, we use satellite imagery, forage quantity data, and three years o...

Ecological Applications, 2011

Conservation Genetics, 2011

In presenting this dissertation in partial fulfillment of the requirements for a doctoral degree ... more In presenting this dissertation in partial fulfillment of the requirements for a doctoral degree at Montana State University, I agree that the Library shall make it available to borrowers under rules of the Library. I further agree that copying of this dissertation is allowable only for scholarly purposes, consistent with "fair use" as prescribed in the U.S. Copyright Law. Requests for extensive copying or reproduction of this dissertation should be referred to ProQuest Information and Learning, 300 North Zeeb Road, Ann Arbor, Michigan 48106, to whom I have granted "the exclusive right to reproduce and distribute my dissertation in and from microform along with the non-exclusive right to reproduce and distribute my

Proceedings of the National Academy of Sciences, 2009

Predators affect prey demography through direct predation and through the costs of antipredator b... more Predators affect prey demography through direct predation and through the costs of antipredator behavioral responses, or risk effects. Experiments have shown that risk effects can comprise a substantial proportion of a predator's total effect on prey dynamics, but we know little about their strength in wild populations, or the physiological mechanisms that mediate them. When wolves are present, elk alter their grouping patterns, vigilance, foraging behavior, habitat selection, and diet. These responses are associated with decreased progesterone levels, decreased calf production, and reduced population size [Creel S, Christianson D, Liley S, Winnie JA (2007) Science 315:960]. Two general mechanisms for the effect of predation risk on reproduction have been proposed: the predation stress hypothesis and the predator-sensitive-food hypothesis. Here, we used enzyme immunoassay to measure fecal glucocorticoid metabolite concentrations for 1,205 samples collected from 4 elk populations...

The risk of predation drives many behavioral responses in prey. However, few studies have directl... more The risk of predation drives many behavioral responses in prey. However, few studies have directly tested whether predation risk alters the way other variables influence prey behavior. Here we use information theory (Akaike's Information Criterion, AICc) in a novel way to test the hypothesis that the decision-making rules governing elk behavior are simplified by the presence of wolves. With elk habitat use as the dependent variable, we test whether the number of independent variables (i.e., the size of the models) that best predict this behavior differ when wolves are present vs absent. Thus, we use AICc scores simply to determine the number of variables to which elk respond when making decisions. We measured habitat use using 2,288 locations from GPS collars on 14 elk, over two winters (14 elk winters), in the Gallatin Canyon portion of the Greater Yellowstone Ecosystem. We found that the use of three major habitat components (grass, conifer, sage) was sensitive to many variables on days that wolves were locally absent, with the best models (ΔAICc≤2) averaging 7.4 parameters. In contrast, habitat use was sensitive to few variables on days when wolves were present: the best models averaged only 2.5 parameters. Because fewer variables affect elk behavior in the presence of wolves, we conclude that elk use simpler decision-making rules in the presence of wolves. This simplification of decisionmaking rules implies that predation risk imposes selection pressures that do not allow prey to respond to other pressures in ways that they otherwise would. If the affected processes are important, then this indirect effect of predation is likely to be important.