Resource Utilization by Foraging Eastern Red Bats (Lasiurus Borealis) in the Ozark Region of Missouri
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The Journal of Wildlife Management 78(3):483–493; 2014; DOI: 10.1002/jwmg.685 Habitat Relations Resource Utilization by Foraging Eastern Red Bats (Lasiurus borealis) in the Ozark Region of Missouri SYBILL K. AMELON,1 Northern Research Station, U.S.D.A. Forest Service, 202 Natural Resource Building, Columbia, MO 65211, USA FRANK R. THOMPSON III, Northern Research Station, U.S.D.A. Forest Service, 202 Natural Resource Building, Columbia, MO 65211, USA JOSHUA J. MILLSPAUGH, University of Missouri, 302 Natural Resource Building, Columbia, MO 65211, USA ABSTRACT Resource selection by animals influences ecological processes such as dispersal, reproduction, foraging, and migration. Little information exists regarding foraging resource selection by bats during the maternity season. We evaluated support for effects of landcover type, landform, and landscape pattern on resource selection by individual foraging female eastern red bats (Lasiurus borealis) during the maternity period and compared resource utilization for all individuals pooled (population level), individuals grouped by geographic location, and individuals grouped by stage of lactation (early, mid, and late). We used a resource utilization function (RUF) to relate landcover and landscape attributes to the utilization distributions of individual bats estimated by the fixed-kernel method. We radio-tracked 64 lactating red bats and estimated utilization distributions for 52 individuals. Mean home range size ranged from 1,041 to 1,588 ha from late to mid lactation. The global RUF model was significantly better than the null RUF model for 36 (70%) individuals and the magnitude and direction of coefficients varied among individuals. Resource utilization at the population level was, on average, positively related to ridges and upland drainage landforms, water landcover, and road density; and negatively related to urban and nonforest landcover and distance to edge. Resource use differed between geographic areas by canopy cover, water landcover, and road density. Canopy closure was positively related to RUF in both areas but was greater in the south area. Percent water negatively related to RUF in the north area and positively related in the south area. Road density had a positive relationship in the north and negative relationship in the south with RUF. We did not find a difference in mean RUF coefficients among lactation groups except for canopy cover. On average, canopy cover had a positive effect on use by bats during early and mid-lactation but a negative effect during late lactation. We suggest that management for red bats consider landscape components and provide a range of composition and structural diversity to enhance foraging use by red bats. In highly forested landscapes, gaps or openings may provide forest edges that are important for foraging and commuting. Published 2014. This article is a U.S. Government work and is in the public domain in the USA. KEY WORDS eastern red bat, foraging, Lasiurus borealis, resource utilization distribution. Insectivorous bats play key functional roles as predators of 2013), has resulted in large declines in hibernating species in insects in temperate ecosystems and declining populations the eastern United States and Canada (Langwig et al. 2012). of this diverse mammalian group have been a concern of Effective management for bat conservation is dependent on resource management and conservation agencies worldwide for our understanding associations between bat populations and several decades (Kunz and Racey 1998, Hutson et al. 2001, their use of resources. Resource selection is a multilevel, Racey and Entwistle 2003). Bat numbers are declining because hierarchical behavioral process used by a species to obtain, of direct persecution, habitat loss, roost disturbance, environ- directly or indirectly, elements needed to survive, reproduce, mental contaminants (Hutson et al. 2001), wind turbines and persist (Johnson 1980, Senft et al. 1987). Studies of (Horn et al. 2008), and climate change (Rebelo et al. 2010). resource selection generally assume that high quality Since 2006, a disease epidemic, white-nose syndrome (WNS) resources will be used more than low quality ones, that associated with the fungal pathogen Psuedogymnoascus destruc- availability is not uniform, and that use of a resource may tans (Geomyces) (Gargas et al. 2009, Minnis and Lindner change with availability (Manly et al. 2002). Despite the importance of such information, resource relationships are poorly understood for most bat species. Bats are highly mobile, which allows them access to a wide range Received: 15 February 2013; Accepted: 30 December 2013 of resources. Historically, foraging resource use has been Published: 21 March 2014 viewed as secondary to roost selection in terms of bat 1E-mail: [email protected] conservation (Kunz 1982). Loss or modification of roosting Amelon et al. Resource Utilization by Eastern Red Bats 483 and foraging habitat, or their juxtaposition, may affect Mager and Nelson 2001). Because red bats seem to use a critical life-history parameters, such as birth rate or adult variety of habitats for foraging, the landscape context and/or survivorship. Recent advances in radio-tracking have allowed foraging resources used at different life-history stages may be for significant progress in our understanding of bat–resource important to meet the diet requirements or other fitness relationships (Barclay and Brigham 1996, Fenton 2003), but characteristics. there continues to be a lack of information regarding use Our objectives were 1) to evaluate support for effects of of foraging habitat by bats (Lacki et al. 2007). Factors landcover type, landform, and landscape pattern on resource potentially influencing the areas used by foraging bats use by individual foraging female red bats during the include prey abundance (Barclay 1985), energetic costs maternity period and at a population level when individual associated with flight mechanics (Aldridge and Rautenbach bat RUFs are averaged (Marzluff et al. 2004, Millspaugh 1987, Brigham et al. 1997), ability to detect and locate prey et al. 2006) and 2) to compare resource use among individuals (Simmons and Stein 1980, Griffin and Thompson 1982, grouped by geographic area (north or south), and stage of Neuweiler 1989, Fenton and Griffin 1997), risk of predation lactation (early, mid, and late). We hypothesized that female (Lesin´ski et al. 2009), proximity of water, and level of human red bats would forage in locations that optimize conditions disturbance in the foraging area (Ford et al. 2006). for fast flight, including ridgetops and drainages, and along Resource selection is often determined by comparing used edges (nonforest next to forest) and/or in locations conducive to available resources, but in practice, it is often difficult to to obtaining water or commuting, including along linear assess availability from an animal’s point of view (Boyce features (such as rivers or roads; Salcedo et al. 1995, Saugey et al. 2002). Quantification of availability usually consists of a et al. 1998, Hutchinson and Lacki 1999). We hypothesized priori or a posteriori measure of the abundance of resources that diverse landcover types may increase use by either in a defined area using classification-based methods that creating additional edge or providing a wider variety of place animal locations into distinct habitat categories for resource options (Hutchinson and Lacki 1999, Elmore analysis (Conner et al. 2003, Miller et al. 2003). In contrast, et al. 2005). We also hypothesized foraging area would the resource utilization function (RUF) approach (Marzluff increase through lactation or at least might change from early et al. 2004, Millspaugh et al. 2006) uses a probability density to late lactation because foraging patterns of females would function (Silverman 1986) that quantifies an individual’s or potentially reflect changing needs of pup development. group’s use of space in a continuous and probabilistic way Specifically, that as females are able to leave pups for longer (Kernohan et al. 2001). The probability density function is periods between nursing bouts, they would use resources often a kernel-based utilization distribution (UD) and depicts farther from roosting sites (Barclay 1989). the probability of an animal occurring at each location within its home range as a function of relocation points (White and STUDY AREA Garrott 1990:146). The UD has been used to relate relative space use to resource attributes in a spatially explicit way using We studied red bats in 2 geographic areas within the Salem the RUF approach (Marzluff et al. 2004). Plateau physiographic region within the Ozark Highlands Widespread or abundant bat species, such as eastern red aquatic sub-region of Missouri (Sowa et al. 2005; Fig. 1). bats (Lasiurus borealis), may be the most ecologically and These areas were primarily within the administrative economically important species because of their role in boundaries of the Houston-Rolla and the Eleven Point ecosystem health (Pierson 1998). Eastern red bats (subse- Ranger Districts of the Mark Twain National Forest and quently referred to only as red bats) are a migratory, tree represent a range of management practices that result in dwelling species that roost singly in tree canopies where they savanna, woodland, and forest communities. The southern hang from leaf petioles or small branches (Perry and Thill 2003, Amelon 2007). Red bats are being killed in high numbers at wind turbines across the continent (Kunz et al. 2007, Winhold et al. 2008). By 2020 an estimated 33,000 to 111,000 migratory bats will be killed annually by wind turbines in the Mid-Atlantic Highlands alone (Kunz et al. 2007). An improved understanding of important site and landscape resources may improve conservation efforts for this species. Red bats consume diverse prey species that include Lepidopterans, Coleopterans, Ephemeropterans, and Hymenopterans (Whitaker 1972, Carter et al. 1998, Clare et al. 2009). Red bats have been reported to use open habitats along the edge of forested areas, agricultural fields, roadways (Salcedo et al. 1995, Saugey et al. 1998), above the tree canopy (Barclay et al. 1999), upland and bottomland Figure 1.