Eric M. Schauber

Effects of Joint Space Use and Group Membership on Contact Rates Among White-Tailed Deer

Abstract Establishment and spread of infectious diseases are controlled by the frequency of contacts among hosts. Although managers can estimate transmission coefficients from the relationship between disease prevalence and age or time, they may wish to quantify or compare contact rates before a disease is established or while it is at very low prevalence. Our objectives were to quantify direct and indirect contacts rates among white-tailed deer (Odocoileus virginianus) and to compare these measures of contact rate with simpler measures of joint space use. We deployed Global Positioning System (GPS) collars on 23 deer near Carbondale, Illinois, USA, from 2002 to 2005. We used location data from the GPS collars to measure pairwise rates of direct and indirect contact, based on a range of proximity criteria and time lags, as well as volume of intersection (VI) of kernel utilization distributions. We analyzed contact rates at a given distance criterion and time lag using mixed-model logistic regression. Direct contact rates increased with increasing VI and were higher in autumn–spring than in summer. After accounting for VI, the estimated odds of direct contact during autumn–spring periods were 5.0–22.1-fold greater (depending on the proximity criterion) for pairs of deer in the same social group than for between-group pairs, but for direct contacts during summer the within:between-group odds ratio did not differ significantly from 1. Indirect contact rates also increased with VI, but the effects of both season and pair-type were much smaller than for direct contacts and differed little as the time lag increased from 1–30 days. These results indicate that simple measures of joint space use are insufficient indices of direct contact because group membership can substantially increase contacts at a given level of joint space use. With indirect transmission, however, group membership had a much smaller influence after accounting for VI. Relationships between contact rates and season, VI, and pair-type were generally robust to changes in the proximity criterion defining a contact, and patterns of indirect contacts were affected little by the choice of time lag from 1–30 days. The use of GPS collars provides a framework for testing hypotheses about the form of contact networks among large mammals and comparing potential direct and indirect contact rates across gradients of ecological factors, such as population density or landscape configuration.

Effects of acorn production and mouse abundance on abundance and Borrelia burgdorferi infection prevalence of nymphal Ixodes scapularis ticks.

Risk of exposure to Lyme disease is a function of the local abundance of nymphal Ixodes ticks that are infected with the etiological agent, the spirochete Borrelia burgdorferi. We monitored abundance of white-footed mice (the principal B. burgdorferi reservoir in the eastern and central United States) and acorns (a critical food resource for mice), and Ixodes scapularis ticks, as well as ambient temperature (cumulative growing degree days) and growing season precipitation, in a forested landscape of southeastern New York State from 1994 to 2000. We found that acorn production in autumn strongly influenced abundance of white-footed mice the following summer and that abundance of mice in summer, when larval ticks are active, influenced the abundance of infected nymphs the following year. Consequently, the abundance of infected nymphal ticks can be predicted from acorn production 1.75 years earlier. Monitoring of natural fluctuations in acorn production thus supports results of prior acorn addition experiments that were conducted at small spatial scales. Growing degree days and precipitation either had no significant effect on density of nymphs or marginally increased the explanatory power of models that included acorns or mouse density as independent variables. We conclude that, at our study site in New York, the risk of human exposure to Lyme disease is affected by mouse density in the prior year and by acorn production 2 years previously.

Space Use and Survival of White-Tailed Deer in an Exurban Landscape

Abstract Exurban development is nonmetropolitan, residential development characterized by a human population density and average property size intermediate between suburban and rural areas. Although growth in exurban areas is outpacing that of urban, suburban, or rural landscapes, studies of deer (Odocoileus spp.) ecology in exurban areas are nonexistent. During 2003–2005, we studied space use (i.e., seasonal home-range and core-area size and habitat use relative to human dwellings) and survival of 43 female white-tailed deer (O. virginianus) in an exurban setting near Carbondale, Illinois. Deer had larger home ranges than most suburban deer populations and generally smaller home ranges than rural deer populations. When we analytically controlled for habitat use, deer exhibited a subtle avoidance of human dwellings, especially during the fawning season. The annual survival rate was among the highest reported in the literature at 0.872 (SE = 0.048). Only 5 deer (cause-specific mortality rate = 0.091) were harvested by hunters, indicating major obstacles for wildlife managers when attempting to manage deer in exurban areas using traditional hunter harvest.

WHAT IS THE BEST PREDICTOR OF ANNUAL LYME DISEASE INCIDENCE: WEATHER, MICE, OR ACORNS?

Predicting fluctuations in annual risk of Lyme disease would be useful in focusing public health efforts. However, several competing hypotheses have been proposed that point to weather variables, acorn production, or mouse abundance as important pre- dictors of Lyme disease risk. We compared the ability of acorn production, mouse density, and four relevant weather variables to predict annual Lyme disease incidence (detrended) between 1992 and 2002 for Dutchess County, New York, and seven states in the northeastern United States. Acorn production and mouse abundance measured in Dutchess County were the strongest predictors (r

Spatiotemporal Variation in a Lyme Disease Host and Vector: Black-Legged Ticks on White-Footed Mice

0.78) of Dutchess County Lyme disease incidence, but the increase in mouse abundance from 1991 to 1992 was contrary to a decrease in Lyme disease in the following years. The Palmer Hydrologic Drought Index (PHDI) was a significant positive predictor of Lyme disease incidence two years later for three states (0.58 # r # 0.88), but summer precipitation was generally negatively correlated with Lyme disease incidence the next year (20.79 # r # 0.02). Mean temperatures for the prior winter or summer showed weak or inconsistent correlations with Lyme disease incidence. In four states, no variable was a statistically significant predictor of Lyme disease incidence. Syn- chrony in Lyme disease incidence between pairs of states was not significantly concordant with synchrony in any weather variable that we examined (0.02 # r # 0.21). We found that acorns and mice were strong predictors of Dutchess County Lyme disease incidence, but their predictive power appeared to be weaker spatially. Moreover, evidence was weak for causal relationships between Lyme disease incidence and the weather variables that we tested. Reliable prediction of Lyme disease incidence may require the identification of new predictors or combinations of biotic and abiotic predictors and may be limited to local scales.

Spatial and Temporal Analysis of Contact Rates in Female White-Tailed Deer

We monitored population density of white-footed mice (Peromyscus leucopus), burdens of immature black-legged ticks (Ixodes scapularis) on mice, and infection prevalence of host-seeking ticks on six forest plots in southeastern New York State from 1995 through 1999. Despite densities of mice that fluctuated two orders of magnitude, average larval and nymphal tick burdens per mouse remained remarkably constant. Spatial variability in mouse density and tick burdens was modest. The total number of larval and nymphal ticks that fed on the mouse population each year depended strongly on population density of mice; a steady increase was observed in both mouse density and total tick meals on mice from 1996 through 1999. The result was a steady increase in the infection prevalence of nymphal and adult ticks with the etiological agent of Lyme disease, Borrelia burgdorferi, over this time. We suggest that fluctuations in population density of mice, combined with possible regulation of tick burdens on mice, may influence risk of human exposure to Lyme disease.

USE OF TRACK PLATES TO QUANTIFY PREDATION RISK AT SMALL SPATIAL SCALES

Abstract White-tailed deer (Odocoileus virginianus) are important game mammals and potential reservoirs of diseases of domestic livestock; thus, diseases of deer are of great concern to wildlife managers. Contact, either direct or indirect, is necessary for disease transmission, but we know little about the ecological contexts that promote intrasexual contact among deer. Using pair-wise direct contacts estimated from Global Positioning System collar locations and joint utilization distributions (JUDs), we assessed habitats in which contacts occur to test whether direct contact rates among female white-tailed deer in different social groups differs among land-cover types. We also tested whether contact rates differed among seasons, lunar phases, and times of day. We obtained locations from 27 female deer for periods of 0.5–17 months during 2002–2006. We designated any simultaneous pair of locations for 2 deer <25 m apart as a direct contact. For each season, we used compositional analysis to compare land-cover types where 2 deer had contact to available land-cover weighted by their JUD. We used mixed-model logistic regression to test for effects of season, lunar phase, and time of day on contact rates. Contact rates during the gestation season were greater than expected from random use in forest and grassland cover, whereas contact rates during the fawning period were greater in agricultural fields than in other land-cover types. Contact rates were greatest during the rut and lowest in summer. Diel patterns of contact rates varied with season, and contact rates were elevated during full moon compared to other lunar periods. Both spatial and temporal analyses suggest that contact between female deer in different social groups occurs mainly during feeding, which highlights the potential impact of food distribution and habitat on contact rates among deer. By using methods to associate contacts and land-cover, we have created beneficial tools for more elaborate and detailed studies of disease transmission. Our methods can offer information necessary to develop spatially realistic models of disease transmission in deer.

Quantifying a Dynamic Risk Landscape: Heterogeneous Predator Activity and Implications for Prey Persistence

Abstract Spatial heterogeneity in risk is a critical component of predator-prey interactions. However, at small spatial scales, it is difficult to quantify predation risk without altering it. We used track plates to measure local predation risk created by white-footed mouse (Peromyscus leucopus) foraging activity on oak-forest plots in Millbrook, New York. Live gypsy moth pupae (Lymantria dispar) were placed at 2 heights on trees and monitored for predation. Pupae deployed on trees visited by mice were more likely to be eaten than those on trees not visited. Logistic regression indicated that predation rates on gypsy moth pupae were positively correlated with track activity, indicating that areas of concentrated mouse activity were areas of heightened risk for gypsy moths. Survival of individual oat grains placed on and 50 cm from track plates were not statistically different, indicating that mice exhibited no detectable behavioral reaction toward track plates. We conclude that track plates offer an economical and reliable means of quantifying local risk of attack by terrestrial mammals without substantially altering the spatial distribution of risk.

Translocation of Swamp Rabbits in Southern Illinois

Spatial heterogeneity in predation risk can ameliorate impacts on prey populations, particularly for prey of generalists. Spatially heterogeneous risk implies the existence of refugia, and the spatial scale of those refugia and their persistence over time affect whether prey can avoid predation by aggregating therein. Our objective was to quantify the magnitude, spatial scale, and temporal persistence of heterogeneity in risk of predation by white-footed mice (Peromyscus leucopus), an abundant generalist predator of gypsy moths (Lymantria dispar) and songbirds. We used track plates to measure white-footed mouse activity at > 170 trees in each of three forest plots in upstate New York during summers of 2003-2005. We quantified the mean and coefficient of variation of track activity among trees by fitting the beta-binomial distribution to data from each plot and study period. We measured temporal persistence by disattenuated autocorrelation, and spatial scale by fitting exponential variograms. Mice were much less abundant in 2005 than the other two years, leading to lower overall track activity but higher coefficient of variation among trees. Mouse track activity at individual trees was positively autocorrelated between monthly study periods in 2003 and 2004, and even between the two years, whereas temporal autocorrelation in 2005 was much weaker. Track activity showed positive spatial autocorrelation over lag distances from approximately 30 to > 1000 m. These findings indicate that mouse activity, and hence risk to their prey, varies substantially in space at spatial and temporal scales that appear responsive to mouse population dynamics. The spatial scale and temporal persistence of that variation imply that prey may benefit from returning to, or failing to disperse from, refugia.

Spatial selection and inheritance: applying evolutionary concepts to population dynamics in heterogeneous space.

Abstract Habitat of Sylvilagus aquaticus (swamp rabbits) in Illinois has been reduced and fragmented due to human land use. Translocation may enable swamp rabbits to colonize isolated habitat patches. We live-trapped and translocated 9 male and 8 female swamp rabbits to unoccupied habitat in southern Illinois in January and February 2004. Eight of 17 translocated rabbits died within 7 days after release. However, mortality rates appeared to drop rapidly over time after release. Predators killed at least 10 of 14 rabbits that died. For conserving swamp rabbits, translocation success is limited by poor live-trapping success and high levels of post-release predation. Intense live-trapping along with predator control in release sites may be necessary to make translocation a viable management strategy.