Influence of natal habitat preference on habitat selection during extra-home range movements in a large ungulate.

Natal habitat preference induction (NHPI) occurs when animals exhibit a preference for new habitat that is similar to that which they experienced in their natal environment, potentially leading to post-dispersal success. While the study of NHPI is typically focused on post-settlement home ranges, we investigated how this behavior may manifest during extra-home range movements (EHRMs), both to identify exploratory prospecting behavior and assess how natal habitat cues may influence path selection before settlement. We analyzed GPS collar relocation data collected during 79 EHRMs made by 34 juvenile and subadult white-tailed deer (Odocoileus virginianus) across an agricultural landscape with highly fragmented forests in Illinois, USA. We developed a workflow to measure multidimensional natal habitat dissimilarity for each EHRM relocation and fit step-selection functions to evaluate whether natal habitat similarity explained habitat selection along movement paths. Across seasons, selection for natal habitat similarity was generally weak during excursive movements, but strong during dispersals, indicating that NHPI is manifested in dispersal habitat selection in this study system and bolstering the hypothesis that excursive movements differ functionally from dispersal. Our approach for extending the NHPI hypothesis to behavior during EHRMs can be applied to a variety of taxa and can expand our understanding of how individual behavioral variation and early life experience may shape connectivity and resistance across landscapes.

Asymmetrical intraguild interactions with coyotes, red foxes, and domestic dogs may contribute to competitive exclusion of declining gray foxes.

Species coexistence is governed by availability of resources and intraguild interactions including strategies to reduce ecological overlap. Gray foxes are dietary generalist mesopredators expected to benefit from anthropogenic disturbance, but populations have declined across the midwestern USA, including severe local extirpation rates coinciding with high coyote and domestic dog occurrence and low red fox occurrence. We used data from a large-scale camera trap survey in southern Illinois, USA to quantify intraguild spatial and temporal interactions among the canid guild including domestic dogs. We used a two-species co-occurrence model to make pairwise assessments of conditional occupancy and detection rates. We also estimated temporal activity overlap among species and fit a fixed-effects hierarchical community occupancy model with the four canid species. We partitioned the posterior distributions to compare gray fox occupancy probabilities conditional on estimated state of combinations of other species to assess support for hypothesized interactions. We found no evidence of broadscale avoidance among native canids and conclude that spatial and temporal segregation were limited by ubiquitous human disturbance. Mean guild richness was two canid species at a site and gray fox occupancy was greater when any combination of sympatric canids was also present, setting the stage for competitive exclusion over time. Domestic dogs may amplify competitive interactions by increasing canid guild size to the detriment of gray foxes. Our results suggest that while human activities can benefit some mesopredators, other species such as gray foxes may serve as bellwethers for habitat degradation with trophic downgrading and continued anthropogenic homogenization.

Does landscape connectivity shape local and global social network structure in white-tailed deer?

Intraspecific social behavior can be influenced by both intrinsic and extrinsic factors. While much research has focused on how characteristics of individuals influence their roles in social networks, we were interested in the role that landscape structure plays in animal sociality at both individual (local) and population (global) levels. We used female white-tailed deer (Odocoileus virginianus) in Illinois, USA, to investigate the potential effect of landscape on social network structure by weighting the edges of seasonal social networks with association rate (based on proximity inferred from GPS collar data). At the local level, we found that sociality among female deer in neighboring social groups (n = 36) was mainly explained by their home range overlap, with two exceptions: 1) during fawning in an area of mixed forest and grassland, deer whose home ranges had low forest connectivity were more social than expected; and 2) during the rut in an area of intensive agriculture, deer inhabiting home ranges with high amount and connectedness of agriculture were more social than expected. At the global scale, we found that deer populations (n = 7) in areas with highly connected forest-agriculture edge, a high proportion of agriculture, and a low proportion of forest tended to have higher weighted network closeness, although low sample size precluded statistical significance. This result implies that infectious disease could spread faster in deer populations inhabiting such landscapes. Our work advances the general understanding of animal social networks, demonstrating how landscape features can underlie differences in social behavior both within and among wildlife social networks.

Relative Preference and Localized Food Affect Predator Space Use and Consumption of Incidental Prey.

Abundant, localized foods can concentrate predators and their foraging efforts, thus altering both the spatial distribution of predation risk and predator preferences for prey that are encountered incidentally. However, few investigations have quantified the spatial scale over which localized foods affect predator foraging behavior and consumption of incidental prey. In spring 2010, we experimentally tested how point-source foods altered how generalist predators (white-footed mice, Peromyscus leucopus) utilized space and depredated two incidental prey items: almonds (Prunus dulcis; highly profitable) and maple seeds (Acer saccharum; less profitable). We estimated mouse population densities with trapping webs, quantified mouse consumption rates of these incidental prey items, and measured local mouse activity with track plates. We predicted that 1) mouse activity would be elevated near full feeders, but depressed at intermediate distances from the feeder, 2) consumption of both incidental prey would be high near feeders providing less-preferred food and, 3) consumption of incidental prey would be contingent on predator preference for prey relative to feeders providing more-preferred food. Mouse densities increased significantly from pre- to post-experiment. Mean mouse activity was unexpectedly greatest in control treatments, particularly <15 m from the control (empty) feeder. Feeders with highly preferred food (sunflower seeds) created localized refuges for incidental prey at intermediate distances (15 to 25m) from the feeder. Feeders with less-preferred food (corn) generated localized high risk for highly preferred almonds <10 m of the feeder. Our findings highlight the contingent but predictable effects of locally abundant food on risk experienced by incidental prey, which can be positive or negative depending on both spatial proximity and relative preference.

Orchid-pollinator interactions and potential vulnerability to biological invasion.

Mutualistic relationships between plants and their pollinators have played a major role in the evolution of biodiversity. While the vulnerability of these relationships to environmental change is a major concern, studies often lack a framework for predicting impacts from emerging threats (e.g. biological invasions). The objective of this study was to determine the reliance of Platanthera ciliaris (orange-fringed orchid) on Papilio palamedes (Palamedes swallowtail butterfly) for pollination and the relative availability of alternative pollinators. Recent declines of P. palamedes larval host plants due to laurel wilt disease (LWD) could endanger P. ciliaris populations that rely heavily on this butterfly for pollination. We monitored pollinator visitation and fruit set and measured nectar spur lengths of P. ciliaris flowers and proboscis lengths of its floral visitors in Jackson County, MS, USA. Papilio palamedes was the primary visitor with minimal visitation by Phoebis sennae (cloudless sulfur butterfly). Lengths of P. ciliaris nectar spurs were similar to proboscis lengths of both pollinator species. Fruit set was moderate with access to pollinators (55 ± 10.8 %), yet failed (0 %) when pollinators were excluded. Visitation increased with inflorescence size, but there was no such pattern in fruit set, indicating that fruit set was not limited by pollinator visitation within the range of visitation rates we observed. Our results are supported by historical data that suggest P. palamedes and P. sennae are important pollinators of P. ciliaris. Although P. sennae may provide supplemental pollination service, this is likely constrained by habitat preferences that do not always overlap with those of P. cilaris. Observed declines of P. palamedes due to LWD could severely limit the reproductive success and persistence of P. ciliaris and similar orchid species populations. This empirical-based prediction is among the first to document exotic forest pests and pathogens as an indirect threat to plant-pollinator interactions.

Social affiliation and contact patterns among white-tailed deer in disparate landscapes: implications for disease transmission.

In social species, individuals contact members of the same group much more often than those of other groups, particularly for contacts that could directly transmit disease agents. This disparity in contact rates violates the assumptions of simple disease models, hinders disease spread between groups, and could decouple disease transmission from population density. Social behavior of white-tailed deer has important implications for the long-term dynamics and impact of diseases such as bovine tuberculosis and chronic wasting disease (CWD), so expanding our understanding of their social system is important. White-tailed deer form matrilineal groups, which inhabit stable home ranges that overlap somewhat with others-a pattern intermediate between mass-action and strict territoriality. To quantify how group membership affects their contact rates and document the spectrum of social affiliation, we analyzed location data from global positioning system (GPS) collars on female and juvenile white-tailed deer in 2 study areas: near Carbondale in forest-dominated southern Illinois (2002-2006) and near Lake Shelbyville in agriculture-dominated central Illinois (2006-2009). For each deer dyad (i.e., 2 individual deer with sufficient overlapping GPS data), we measured space-use overlap, correlation of movements, direct contact rate (simultaneous GPS locations < 10 m apart), and indirect contact rate (GPS locations < 10 m apart when offset by 1 or 3 days). Direct contact rates were substantially higher for within-group dyads than between-group dyads, but group membership had little apparent effect on indirect contact rates. The group membership effect on direct contact rates was strongest in winter and weakest in summer, with no apparent difference between study areas. Social affiliations were not dichotomous, with some deer dyads showing loose but positive affiliation. Even for obvious within-group dyads, their strength of affiliation fluctuated between years, seasons, and even days. Our findings highlight the poor fit between deer behavior and simple models of disease transmission and, combined with previous infection data, suggest that direct contact is the primary driver of CWD transmission among free-living female and juvenile white-tailed deer.

Familiarity breeds contempt: combining proximity loggers and GPS reveals female white-tailed deer (Odocoileus virginianus) avoiding close contact with neighbors.

Social interactions can influence infectious disease dynamics, particularly for directly transmitted pathogens. Therefore, reliable information on contact frequency within and among groups can better inform disease modeling and management. We compared three methods of assessing contact patterns: (1) space-use overlap (volume of interaction [VI]), (2) direct contact rates measured by simultaneous global positioning system (GPS) locations (<10 m apart), and (3) direct contact rates measured by proximity loggers (PLs; 1-m detection) among female white-tailed deer (Odocoileus virginianus). We calculated the PL∶GPS contact ratios to see whether both devices reveal similar contact patterns and thus predict similar pathogen transmission patterns. Contact rates measured by GPS and PLs were similarly high for two within-group dyads (pairs of deer in the same social groups). Dyads representing separate but neighboring groups (high VI) had PL∶GPS contact ratios near zero, whereas dyads further apart (intermediate VI) had higher PL∶GPS contact ratios. Social networks based on PL contacts showed the fewest connected individuals and lowest mean centrality measures; network metrics were intermediate when based on GPS contacts and greatest when based on VI. Thus, the VI network portrayed animals to be more uniformly and strongly connected than did the PL network. We conclude that simultaneous GPS locations, compared with PLs, substantially underestimate the impact of group membership on direct contact rates of female deer and make networks appear more connected. We also present evidence that deer coming within the general vicinity of each other are less likely to come in close contact if they are in neighboring social groups than deer whose home ranges overlap little if at all. Combined, these results provide evidence that direct transmission of disease agents among female and juvenile white-tailed deer is likely to be constrained both spatially and by social structure, more so than GPS data alone would suggest.

Quantifying a dynamic risk landscape: heterogeneous predator activity and implications for prey persistence.

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.

Organisms in highly suitable sites generally produce more offspring, and offspring can inherit this suitability by not dispersing far. This combination of spatial selection and spatial inheritance acts to bias the distribution of organisms toward suitable sites and thereby increase mean fitness (i.e., per capita population increase). Thus, population growth rates in heterogeneous space change over time by a process conceptually analogous to evolution by natural selection, opening avenues for theoretical cross-pollination between evolutionary biology and ecology. We operationally define spatial inheritance and spatial selective differential and then combine these two factors in a modification of the breeder's equation, derived from simple models of population growth in heterogeneous space. The modified breeder's equation yields a conservative criterion for persistence in hostile environments estimable from field measurements. We apply this framework for understanding gypsy moth population persistence amidst abundant predators and find that the predictions of the modified breeder's equation match initial changes in population growth rate in independent simulation output. The analogy between spatial dynamics and natural selection conceptually links ecology and evolution, provides a spatially implicit framework for modeling spatial population dynamics, and represents an important null model for studying habitat selection.