Southern Illinois University Carbondale OpenSIUC Publications Department of Zoology 5-2015 Spatial and Temporal Structure of a Mesocarnivore Guild in Midwestern North America Damon B. Lesmeister Southern Illinois University Carbondale Clayton K. Nielsen Southern Illinois University Carbondale Eric M. Schauber Southern Illinois University Carbondale, [email protected] Eric Hellgren Southern Illinois University Carbondale Follow this and additional works at: http://opensiuc.lib.siu.edu/zool_pubs Recommended Citation Lesmeister, Damon B., Nielsen, Clayton K., Schauber, Eric M. and Hellgren, Eric. "Spatial and Temporal Structure of a Mesocarnivore Guild in Midwestern North America." Wildlife Monographs 191 (May 2015): 1-61. doi:10.1002/wmon.1015. This Article is brought to you for free and open access by the Department of Zoology at OpenSIUC. It has been accepted for inclusion in Publications by an authorized administrator of OpenSIUC. For more information, please contact [email protected]. This is the pre-peer reviewed version of: Lesmeister, D. B., C. K. Nielsen, E. M. Schauber, and E. C. Hellgren. 2015. Spatial and temporal structure of a mesocarnivore guild in midwestern North America. Wildlife Monographs , 191:1-61. Which has been published in final form at: DOI: 10:1002/wmon.1015. Lesmeister et al. • Factors Influencing Carnivore Guild Structure 1 ACKNOWLEDGMENTS (FUNDING) Funding for this research came from the Illinois Department of Natural Resources, Federal Aid Project W-135-R. Additional funding was provided by the Cooperative Wildlife Research Laboratory, Department of Zoology, and the Graduate School at Southern Illinois University Carbondale. For covering publication costs, we thank the Illinois Department of Natural Resources. Lesmeister et al. • Factors Influencing Carnivore Guild Structure 2 20 April 2013 Damon B. Lesmeister Crab Orchard National Wildlife Refuge United States Fish and Wildlife Service 8588 Rt. 148 Marion, IL 62959 Tel: 618-998-5917 Fax: 618-997-8961 Email: [email protected] RH: Lesmeister et al. • Factors Influencing Carnivore Guild Structure Factors Influencing the Spatial and Temporal Structure of a Carnivore Guild in the Central Hardwood Region DAMON B. LESMEISTER 12, Cooperative Wildlife Research Laboratory, Southern Illinois University Carbondale, 251 Life Science II, Carbondale, IL 62901, USA. ERIC C. HELLGREN, Cooperative Wildlife Research Laboratory, Southern Illinois University Carbondale, 251 Life Science II, Carbondale, IL 62901, USA. CLAYTON K. NIELSEN, Cooperative Wildlife Research Laboratory, Southern Illinois University Carbondale, 251 Life Science II, Carbondale, IL 62901, USA. ERIC M. SCHAUBER, Cooperative Wildlife Research Laboratory, Southern Illinois University Carbondale, 251 Life Science II, Carbondale, IL 62901, USA. ABSTRACT Ecological communities are most commonly structured by a mixture of bottom-up processes such as habitat or prey, competition within the same trophic level, and top-down forces from higher trophic levels. Carnivore guilds play a vital role in the broader ecological community by stabilizing or destabilizing food webs. Consequently, factors influencing the structure of carnivore guilds can be critical to ecosystem patterns. Coyotes ( Canis latrans ), 1 Present address: Crab Orchard National Wildlife Refuge, United States Fish and Wildlife Service, 8588 Route 148, Marion, IL 62959, USA. 2Email: [email protected] Lesmeister et al. • Factors Influencing Carnivore Guild Structure 3 bobcats ( Lynx rufus ), gray foxes ( Urocyon cinereoargenteus ), raccoons ( Procyon lotor ), red foxes ( Vulpes vulpes ), and striped skunks ( Mephitis mephitis ) occur sympatrically throughout much of their geographic ranges in North America and overlap in resource use, indicating potential for interspecific interactions. Although much is known about space use, habitat relationships, and activity patterns of the individual species separately, little is known about factors that facilitate coexistence and how interactions within this guild influence distribution, activity, and survival of the smaller carnivores. For example, gray fox populations appear to have declined in Illinois since the early 1990s and it is unknown if the increase in bobcat and coyote populations during the same time period is the cause. We conducted a large-scale non- invasive carnivore survey using an occupancy modeling framework to quantify factors affecting the structure of this widely-occurring carnivore guild. We used baited remote cameras during 3- week surveys to detect carnivores at 1,118 camera-points in 357 2.6-km 2 sections (clusters of 3– 4 cameras/section) in the 16 southernmost counties of Illinois (16,058 km 2) during January– April, 2008–2010. We collected microhabitat data at each camera-point and landscape-level habitat data for each camera-cluster. In a multi-stage approach, we used information-theoretic methods to develop and evaluate models for detection, species-specific habitat occupancy, multi- species co-occupancy, and multi-season (colonization and extinction) occupancy dynamics. We developed hypotheses for each species regarding the occupancy of areas based on anthropogenic features, prey availability, landscape complexity, and vegetative landcover. We used photographic data, Poisson regression, and mixed-model logistic regression to quantify temporal activity of carnivores in the study area and how interspecific factors influence temporal patterns of activity. Lesmeister et al. • Factors Influencing Carnivore Guild Structure 4 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 at a camera point decreased after a bobcat photograph was recorded, suggesting avoidance of remote cameras. Across southern Illinois, bobcat occupancy at the camera-point and camera-cluster scale ( ψ̂ point = 0.24 ± 0.04, camera-cluster ψ̂ cluster = 0.75 ± 0.06) was negatively influenced by anthropogenic features and infrastructure. Bobcats had high rates of colonization ( γ̂ = 0.86) and low rates of extinction ( ε̂ = 0.07) during the study, suggesting an expanding population, but agricultural land was less likely to be colonized. Nearly all camera clusters were occupied by coyotes ( ψ̂ cluster = 0.95 ± 0.03) during the entire study. At the camera- point scale, coyote occupancy (overall ψ̂ point = 0.58 ± 0.03) was higher in hardwood forest stands with open understories than in other habitats. Compared to coyotes, gray foxed occupied a lower portion of the study area (ψ̂ point = 0.13 ± 0.01, ψ̂ cluster = 0.29 ± 0.03) at all scales. At the camera-cluster scale, gray fox occupancy was highest in spatially-complex areas with high proportions of forest, and positively related to anthropogenic features within 100% estimated home-range buffers. Red foxes occupied a similar proportion of the study area as gray foxes ( ψ̂ point = 0.12 ± 0.02, ψ̂ cluster = 0.26 ± 0.04), but were more closely associated with anthropogenic features. Indeed, at all three scales of red fox occupancy analysis, only anthropogenic feature models occurred in the 90% confidence set. Camera-cluster extinction probabilities were higher for both gray foxes ( ε̂ = 0.57) and red foxes (ε̂ = 0.35) than their colonization rates (gray fox γ̂ = 0.16, red fox γ̂ = 0.06), suggesting both species may be declining in southern Illinois. Striped skunks occupied a large portion of the Lesmeister et al. • Factors Influencing Carnivore Guild Structure 5 study area ( ψ̂ point = 0.47 ± 0.01, ψ̂ cluster = 0.79 ± 0.03) and were associated primarily with anthropogenic features, especially if the features were surrounded by agricultural land and not forest. Raccoons were essentially ubiquitous within the study area, being photographed in 99% of camera clusters. In some instances, the presence of other carnivores appeared to be an important factor in the occupancy of the 4 smaller species, but in general, habitat models were more supported than co-occurrence models. Habitat had a stronger influence on the occupancy of gray foxes and red foxes than did the presence of bobcats. However, the level of red fox activity was negatively correlated with bobcat activity. Gray fox occupancy and level of activity were reduced in camera-clusters occupied by coyotes, but were not related to bobcat occupancy. When not considering the presence of coyotes, gray foxes appeared to use camera points with fewer hardwood and more conifer trees, which was counter to previous findings. However, when adding the effect of coyote presence, gray fox ψpoint models indicated a positive relationship with hardwood stands. Therefore, gray foxes were more likely to occupy camera points in hardwood stands than conifer stands if coyotes were also present; suggesting that hardwood stands may enhance gray fox-coyote coexistence. The 2 fox species appeared to co-occur with each other at the camera-point scale more frequently than expected on the basis of their individual selection of habitat. Similarly, camera- point occupancy of red foxes was higher when coyotes were present. These apparent canid associations may be a response to locally-high prey abundance or an unmeasured habitat variable. Activity levels of raccoons, bobcats, and coyotes