Spatial and temporal structure of a mesocarnivore guild in midwestern north America Estructura Espacial y Temporal de un Gremio Mesocarnívoro en el medio oeste de América del Norte Structure Spatio-Temporelle d'une Guilde de Mésocarnivores dans le Midwest Nord Américain

Abstract

Carnivore guilds play a vital role in ecological communities by cascading trophic effects, energy and nutrient transfer, and stabilizing or destabilizing food webs. Consequently, the structure of carnivore guilds can be critical to ecosystem patterns. Body size is a crucial influence on intraguild interactions, because it affects access to prey resources, effectiveness in scramble competition, and vulnerability to intraguild predation. Coyotes (Canis latrans), bobcats (Lynx rufus), gray foxes (Urocyon cinereoargenteus), raccoons (Procyon lotor), red foxes (Vulpes vulpes), and striped skunks (Mephitis mephitis) occur sympatrically throughout much of North America and overlap in resource use, indicating potential for interspecific interactions. Although much is known about the autecology of the individual species separately, little is known about factors that facilitate coexistence and how interactions within this guild influence distribution, habitat use, and temporal activity of the smaller carnivores. To assess how habitat autecology and interspecific interactions affect the structure of this widespread carnivore guild, we conducted a large-scale, non-invasive carnivore survey using an occupancy modeling framework. We deployed remote cameras during 3-week surveys to detect carnivores at 1,118 camera locations in 357 2.6-km² sections (3–4 cameras/section composing a cluster) in the 16 southernmost counties of Illinois (16,058 km²) during January–April, 2008–2010. We characterized microhabitat at each camera location and landscape-level habitat features for each camera cluster. In a multistage approach, we used information-theoretic methods to evaluate competing models for detection, species-specific habitat occupancy, multispecies co-occupancy, and multiseason (colonization and extinction) occupancy dynamics. We developed occupancy models for each species to represent hypothesized effects of anthropogenic features, prey availability, landscape complexity, and vegetative land cover. We quantified temporal activity patterns of each carnivore species based on their frequency of appearance in photographs. Further, we assessed whether smaller carnivores shifted their diel activity patterns in response to the presence of potential competitors. Of the 102,711 photographs of endothermic animals, we recorded photographs of bobcats (n = 412 photographs), coyotes (n = 1,397), gray foxes (n = 546), raccoons (n = 40,029), red foxes (n = 149), and striped skunks (n = 2,467). Bobcats were active primarily during crepuscular periods, and their activity was reduced with precipitation and higher temperatures. The probability of detecting bobcats decreased after a bobcat photograph was recorded, suggesting avoidance of remote cameras after the first encounter. Across southern Illinois, bobcat occupancy at the camera-location and camera-cluster scale ( = 0.24 ± 0.04, camera cluster  = 0.75 ± 0.06) was negatively influenced by anthropogenic features and infrastructure. Bobcats had high rates of colonization ( = 0.86) and low rates of extinction ( = 0.07), suggesting an expanding population, but agricultural land was less likely to be colonized. Nearly all camera clusters were occupied by coyotes ( = 0.95 ± 0.03). At the local scale, coyote occupancy ( = 0.58 ± 0.03) was higher in hardwood forest stands with open understories than in other areas. Compared to coyotes, gray foxes occupied a smaller portion of the study area ( = 0.13 ± 0.01,  = 0.29 ± 0.03) at all scales. At the scale of the camera cluster, gray fox occupancy was highest in fragmented areas with high proportions of forest, and positively related to anthropogenic features within 100% home-range buffers. Red foxes occupied a similar proportion of the study area as gray foxes ( = 0.12 ± 0.02,  = 0.26 ± 0.04) but were more closely associated with anthropogenic features. Only anthropogenic feature models made up the 90% confidence set at all scales of analysis for red foxes. Extinction probabilities at the scale of the camera cluster were higher for both gray foxes ( = 0.57) and red foxes ( = 0.35) than their colonization rates (gray fox  = 0.16, red fox  = 0.06), suggesting both species may be declining in southern Illinois. Striped skunks occupied a large portion of the study area ( = 0.47 ± 0.01,  = 0.79 ± 0.03) and were associated primarily with anthropogenic features. Raccoons were essentially ubiquitous within the study area, being photographed in 99% of camera clusters. We observed little evidence for spatial partitioning based on interspecific interactions, with the exception of the gray fox-coyote pairs, and found that habitat preferences were more important in structuring the carnivore community. Habitat had a stronger influence on the occupancy of foxes than did the presence of bobcats. However, the level of red fox activity was negatively correlated with bobcat activity. Gray fox occupancy and the number of detections within occupied sites were reduced in camera clusters occupied by coyotes but not bobcat occupancy. Overall, gray fox occupancy was highest at camera locations with fewer hardwood and more conifer trees. However, gray foxes were more likely to occupy camera locations in hardwood stands than conifer stands if coyotes were also present indicating that hardwood stands may enhance gray fox-coyote coexistence. The 2 fox species appeared to co-occur with each other at the local scale more frequently than expected based on their individual selection of habitat. Similarly, occupancy of camera location by red foxes was higher when coyotes were present. These positive spatial associations among canids may be a response to locally high prey abundance or unmeasured habitat variables. Activity levels of raccoons, bobcats, and coyotes were all positively correlated. Overall, our co- occurrence and activity models indicate competitor-driven adjustments in space use among members of a carnivore community might be the exception rather than the norm. Nevertheless, although our results indicate that gray foxes and red foxes currently coexist with bobcats and coyotes, their distribution appears to be contracting on our study area. Coexistence of foxes with larger carnivores may be enhanced by temporal partitioning of activity and by habitat features that reduce vulnerability of intraguild predation. For instance, hardwood stands may contain trees with structure that enhances tree-climbing by gray foxes, a behavior that probably facilitates coexistence with coyotes. Efforts to enhance gray fox populations would likely benefit from increasing the amount of mature oak-hickory forest. Additionally, the varying results from different scales of analyses underscore the importance of considering multiple spatial scales in carnivore community studies. © 2015 The Wildlife Society.