HABITAT ASSOCIATIONS AMONG BATS ON REDSTONE ARSENAL, ALABAMA Except where reference is made to the work of others, the work described in this thesis is my own or was done in collaboration with my advisory committee. This thesis does not include proprietary or classified information. _____________________________________ Sara Elizabeth Gardner Certificate of Approval: _________________________ _________________________ James B. Armstrong Troy L. Best, Chairman Professor Professor Forestry and Wildlife Sciences Biological Sciences _________________________ _________________________ Robert S. Lishak George T. Flowers Associate Professor Dean Biological Sciences Graduate School HABITAT ASSOCIATIONS AMONG BATS ON REDSTONE ARSENAL, ALABAMA Sara Elizabeth Gardner A Thesis Submitted to the Graduate Faculty of Auburn University in Partial Fulfillment of the Requirements for the Degree of Master of Science Auburn, Alabama December 19, 2008 iii HABITAT ASSOCIATIONS AMONG BATS ON REDSTONE ARSENAL, ALABAMA Sara Elizabeth Gardner Permission is granted to Auburn University to make copies of this thesis at its discretion, upon request of individuals or institutions and at their expense. The author reserves all publication rights. ______________________________ Signature of Author ______________________________ Date of Graduation iv VITA Sara Elizabeth Gardner, daughter of Gary Thomas and Ruth Ann Gardner, was born 6 July 1981 in Atlanta, Georgia. In 1999, she graduated magna cum laude from Cleveland High School in Cleveland, Tennessee. In 2003, she graduated from Middle Tennessee State University in Murfreesboro, Tennessee, with a Bachelor of Science degree in Biology with a concentration in plant biology. After working as a Research Science Technician for the Missouri Department of Conservation for 2 years, she decided to further her academic education. She entered graduate school at Auburn University in August 2005, where she worked as a Graduate Teaching Assistant and Graduate Research Assistant in the Department of Biological Sciences during her graduate program. v THESIS ABSTRACT HABITAT ASSOCIATIONS AMONG BATS ON REDSTONE ARSENAL, ALABAMA Sara E. Gardner Master of Science, December 19, 2008 (B.S., Middle Tennessee State University, 2003) 108 Typed Pages Directed by Troy L. Best As forests become more fragmented, more research about use of forests by bats at the landscape level is needed. I employed 2 methods to investigate use of forest by foraging bats; mist-net surveys and ultrasonic-detection surveys. I sampled at a total of 341 sites (248 mist-net sites and 93 ultrasonic-detection sites). I captured 185 bats in mist nets at 82 sites representing 7 species, and I ultrasonically detected bats at 45 sites representing 6 species. This study documented use of Redstone Arsenal as foraging habitat for one endangered species (Myotis grisescens) and two species of highest conservation concern (M. austroriparius and M. septentrionalis). I also compared number of species detected per night using the 2 methods. The ultrasonic-detection method detected more species per night. Unlike other studies, I detected more species overall using mist-net surveys (7 species) than ultrasonic detection (6 species). All species that I recorded using ultrasonic-detection were captured in mist nets. vi ACKNOWLEDGMENTS I thank my M.S. committee, Troy L. Best, James B. Armstrong, and Robert S. Lishak, for comments and suggestions; Lisa A. McWilliams, Charles H. Kilgore, Amber C. Dunn, Victoria Antoniak, Christina Willis, and the 2006 and 2007 Field Biology and Ecology classes for assistance in the field; David Nixon, Robert Richey (a.k.a. Legs), Danny Dunn, and Gabrielle Ehinger of the Natural Resources Department at Redstone Arsenal for providing administrative support and assistance in the field, Eric Britzke for help with analysis of bat calls, Mark MacKenzie for assistance with GIS, Kim Livengood and Chris Corben for advice and instruction involving AnaBat, Michael C. Wooten and David M. Shannon for advice with statistical analyses, and the Best Lab and friends, Kyle Barrett, L. Michelle Gilley, Samuel J. Hirt, Vikki Peterson, and Jeremy A. White. This study was funded primarily by Redstone Arsenal; additional support was provided by Auburn University, Department of Biological Sciences. Thanks to Steve Samoray for help in the field, suggestions, and support. vii Style manual or journal used: Journal of Mammalogy Computer software used: Microsoft Word, Microsoft Excel, Microsoft Access, MINITAB 14, SAS 9.1, ArcGIS 9.1, Analook v. 4.9j viii TABLE OF CONTENTS PAGE LIST OF TABLES.............................................................................................................. x LIST OF FIGURES ........................................................................................................ xiii CHAPTER 1 HABITAT ASSOCIATIONS OF BATS ON REDSTONE ARSENAL, MADISON CO., ALABAMA, AS DETERMINED BY MIST-NET SURVEYS............ 1 ABSTRACT....................................................................................................................... 1 INTRODUCTION ............................................................................................................. 1 MATERIALS AND METHODS....................................................................................... 3 RESULTS .......................................................................................................................... 5 DISCUSSION.................................................................................................................. 11 LITERATURE CITED .................................................................................................... 15 CHAPTER 2: HABITAT ASSOCIATIONS AMONG BATS ON REDSTONE ARSENAL, MADISON CO., ALABAMA, AS DETERMINED BY ULTRASONIC-DETECTION SURVEYS.................................................................35 ABSTRACT.......................................................................................................................35 INTRODUCTION ........................................................................................................... 35 MATERIALS AND METHODS..................................................................................... 37 RESULTS ........................................................................................................................ 38 DISCUSSION.................................................................................................................. 43 ix LITERATURE CITED .................................................................................................... 46 CHAPTER 3: COMPARING SURVEY METHODS TO ELUCIDATE HABITAT ASSOCIATIONS OF BATS ON REDSTONE ARSENAL, ALABAMA.......................62 ABSTRACT..................................................................................................................... 62 INTRODUCTION ........................................................................................................... 62 MATERIALS AND METHODS..................................................................................... 63 RESULTS ........................................................................................................................ 63 DISCUSSION.................................................................................................................. 64 LITERATURE CITED .................................................................................................... 65 APPENDIX 1................................................................................................................... 69 APPENDIX 2................................................................................................................... 81 APPENDIX 3................................................................................................................... 87 x LIST OF TABLES PAGE Table 1.1. Species, sample size (n), sex, age-class, and reproductive status of bats captured at Redstone Arsenal, Madison Co., Alabama: YOY, young-of- the-year; NN, not pregnant and not lactating; L, lactating; PL, post- lactating; PR, pregnant; I, immature; S, scrotal; NS, non-scrotal; U, undetermined ..................................................................................................22 Table 1.2. Significant habitat associations of bats on Redstone Arsenal, Madison Co., Alabama, 2005-2007, as determined by GLM comparing mean area of each habitat type where a species was captured versus sites where it was not captured. A positive sign indicates habitats where bats were captured and a negative sign indicates habitat where bats were not captured. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01 .....................23 Table 1.3. Significant habitat associations for bats on Redstone Arsenal, Madison Co., Alabama, 2005-2007, as determined by GLM comparing mean area of each habitat type at sites where a species was captured against random sites. A positive sign indicates habitats where bats were detected and a negative sign indicates habitats where bats were not detected. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. ....................24 xi Table 1.4. Species of bats that were not captured at Redstone Arsenal, Madison Co., Alabama, but potentially occur there..............................................................25 Table 2.1. Significant habitat associations for bats that occur on Redstone Arsenal, Madison Co., Alabama, 2005-2007, as determined by GLM comparing mean area of each habitat type at sites where a species was detected against sites where that species was not detected. A positive sign indicates habitats where bats were detected and a negative sign indicates habitat where bats were not detected. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01 ..............................................................................49 Table 2.2. Significant habitat associations for bats that occur on Redstone Arsenal, Madison Co., Alabama, 2005-2007, as determined by GLM comparing mean area of each habitat type at sites where a species was detected against random sites. A positive sign indicates habitats where bats were detected and a negative sign indicates habitat where bats were not detected. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01 .................................................................................................................50 Table 2.3. Species of bats that occur in the geographic range of Redstone Arsenal, Madison Co., Alabama, that were not detected during this study ..................51 xii Table 3.1. Values used in a t-test to compare number of species per night that were captured in mist nets and by AnaBat on Redstone Arsenal, Madison Co., Alabama..........................................................................................................68 xiii LIST OF FIGURES Figure 1.1 Sites where mist nets were placed to capture bats on Redstone Arsenal, Madison, Co., Alabama, summers 2005-2007 ...............................................26 Figure 1.2 Dates bats were captured at Redstone Arsenal, Madison Co., Alabama. These data indicate that number of bats captured increased as summers progressed during 2005-2007 .........................................................................27 Figure 1.3 Average area (m 2 ) of habitat types at the 250-m spatial scale at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates capture sites and a striped bar indicates no-capture sites for a) Eptesicus fuscus, b) Lasiurus borealis, c) Myotis austroriparius, d) M. grisescens, e) Nycticeius humeralis, and f) Perimyotis subflavus. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01..................................................28 Figure 1.4 Average area (m 2 ) of habitat types at the 500-m spatial scale at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates capture sites and a striped bar indicates no-capture sites for a) Eptesicus fuscus, b) Lasiurus borealis, c) Myotis austroriparius, d) M. grisescens, e) Nycticeius humeralis, and f) Perimyotis subflavus. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01..................................................29 Figure 1.5 Average area (m 2 ) of habitat types at the 1,000-m spatial scale at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates capture xiv sites and a striped bar indicates no-capture sites for a) Eptesicus fuscus, b) Lasiurus borealis, c) Myotis austroriparius, d) M. grisescens, e) Nycticeius humeralis, and f) Perimyotis subflavus. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01..................................................30 Figure 1.6 Average area (m 2 ) of habitat types at the 250-m spatial scale at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates capture sites and a striped bar indicates random sites for a) Eptesicus fuscus, b) Lasiurus borealis, c) Myotis austroriparius, d) M. grisescens, e) Nycticeius humeralis, and f) Perimyotis subflavus. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01..................................................31 Figure 1.7 Average area (m 2 ) of habitat types at the 500-m spatial scale at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates capture sites and a striped bar indicates random sites for a) Eptesicus fuscus, b) Lasiurus borealis, c) Myotis austroriparius, d) M. grisescens, e) Nycticeius humeralis, and f) Perimyotis subflavus. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01..................................................32 Figure 1.8 Average area (m 2 ) of habitat types at the 1,000-m spatial scale at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates capture sites and a striped bar indicates random sites for a) Eptesicus fuscus, b) Lasiurus borealis, c) Myotis austroriparius, d) M. grisescens, e) Nycticeius humeralis, and f) Perimyotis subflavus. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01..................................................33 xv Figure 1.9 Average area (m 2 ) of habitat types at the a) 250-m, b) 500-m, and c) 1,000- m spatial scale at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates sites where nets were placed (captures and no-captures) and a striped bar indicates random sites. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. ...............................................................34 Figure 2.1. Sites were AnaBat detectors were placed on Redstone Arsenal, Madison, Co., Alabama, summer 2007. .........................................................................52 Figure 2.2. Photo of the weather-proof housing in which AnaBat detectors were placed during the study of foraging habitats of bats at Redstone Arsenal, Madison Co., Alabama, 2007. ........................................................................53 Figure 2.3 Representative echolocation calls for a) Eptesicus fuscus, b) Lasiurus borealis, c) Myotis grisescens, d) Nycticeius humeralis, and e) Perimyotis subflavus as detected by an acoustic-monitoring device at Redstone Arsenal, Madison, Co., Alabama....................................................................54 Figure 2.4 Average area (m 2 ) of habitat types at the 250-m spatial scale at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates detection sites and a striped bar indicates no-detection sites for a) Eptesicus fuscus, b) Lasiurus borealis, c) Myotis grisescens, d) Nycticeius humeralis, and e) Perimyotis subflavus. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. ...............................................................55 Figure 2.5 Average area (m 2 ) of habitat types at the 500-m spatial scale at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates detection sites and a striped bar indicates no-detection sites for a) xvi Eptesicus fuscus, b) Lasiurus borealis, c) Myotis grisescens, d) Nycticeius humeralis, and e) Perimyotis subflavus. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. ...............................................................56 Figure 2.6 Average area (m 2 ) of habitat types at the 1,000-m spatial scale at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates detection sites and a striped bar indicates no-detection sites for a) Eptesicus fuscus, b) Lasiurus borealis, c) Myotis grisescens, d) Nycticeius humeralis, and e) Perimyotis subflavus. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. ...............................................................57 Figure 2.7 Average area (m 2 ) of habitat types at the 250-m spatial scale at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates detection sites and a striped bar indicates random sites for a) Eptesicus fuscus, b) Lasiurus borealis, c) Myotis grisescens, d) Nycticeius humeralis, and e) Perimyotis subflavus. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. ...............................................................58 Figure 2.8 Average area (m 2 ) of habitat types at the 500-m spatial scale at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates detection sites and a striped bar indicates random sites for a) Eptesicus fuscus, b) Lasiurus borealis, c) Myotis grisescens, d) Nycticeius humeralis, and e) Perimyotis subflavus. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. ...............................................................59 Figure 2.9 Average area (m 2 ) of habitat types at the 1,000-m spatial scale at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates xvii detection sites and a striped bar indicates random sites for a) Eptesicus fuscus, b) Lasiurus borealis, c) Myotis grisescens, d) Nycticeius humeralis, and e) Perimyotis subflavus. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. ...............................................................60 Figure 2.10 Average area (m 2 ) of habitat types at the a) 250-m, b) 500-m, and c) 1,000-m spatial scales at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates sites where detectors were placed and a striped bar indicates random sites. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. .............................................................................61 1 CHAPTER 1 HABITAT ASSOCIATIONS OF BATS ON REDSTONE ARSENAL, MADISON CO., ALABAMA, AS DETERMINED BY MIST-NET SURVEYS ABSTRACT Little is known about the ecology of forest-dwelling bats, particularly their foraging ecology. I assessed associations of foraging habitats and bats in northern Alabama by capturing bats in mist nets and comparing species of bats captured to types of habitats. Multivariate analysis of variance indicated that the eastern red bat (Lasiurus borealis) was associated with evergreen forest, southeastern myotis (Myotis austroriparius) was associated with open water and wetlands, evening bat (Nycticeius humeralis) was associated with wetlands, and perimyotis (Perimyotis subflavus) was associated with open water. This study provided data for use in creating management plans for foraging habitat among southeastern species of bats. INTRODUCTION Previous studies of the ecology of bats focused on species that use caves or man- made structures. Recently, interest in learning more about ecology of bats has focused on forest-dwelling species and answering questions related to management of forested habitats to promote continued use by bats (Brigham 2007; Miller et al. 2003). Generally, use of landscape in forested habitats by bats for foraging is not known (Kalcounis- R?ppell et al. 2005; Miller et al. 2003), but we do know that biodiversity in forests is 2 being lost due to habitat fragmentation (Fahrig 2003; Gorresen and Willig 2004) and that patches of remnant forest are becoming increasingly important for bats (Evelyn and Stiles 2007). There is not sufficient information to supply land managers with advice about managing forests to assure presence of appropriate foraging habitats for bats (Miller et al. 2003). Miles et al. (2006) contended that more landscape-level investigations of use of habitats by bats are needed. Composition of landscape (Gorresen and Willig 2004) and type of forest (Kalcounis et al. 1999; Patriquin and Barclay 2003) can be used to determine abundance of species. Gorresen et al. (2005) suggested that it is necessary to use a range of scales at which bats function to determine use of landscape because the landscape is used for both roosting and foraging. This information is crucial in developing management plans for species listed as threatened or endangered by federal and state wildlife-management agencies. The bat fauna of Alabama includes 15 species; southeastern myotis (Myotis austroriparius), gray myotis (M. grisescens), little brown myotis (M. lucifugus), northern myotis (M. septentrionalis), Indiana myotis (M. sodalis), perimyotis (Perimyotis subflavus), big brown bat (Eptesicus fuscus), silver-haired bat (Lasionycteris noctivagans), eastern red bat (Lasiurus borealis), hoary bat (L. cinereus), northern yellow bat (L. intermedius), Seminole bat (L. seminolus), evening bat (Nycticeius humeralis), Rafinesque's big eared bat (Corynorhinus rafinesquii), and Brazilian free-tailed bat (Tadarida brasiliensis--modified from Best 2004b; Hall 1981; Harvey et al. 1999). Two of these species are listed as endangered by the United States Fish and Wildlife Service (M. grisescens and M. sodalis), one is a state-listed species of highest conservation concern (C. rafinesquii), five are of high conservation concern (L. intermedius, M. 3 austroriparius, M. lucifugus, M. septentrionalis, and T. brasiliensis), and two are poorly known (L. noctivagans, L. cinereus; Best 2004a). Relatively little is known about the ecology of bats in Alabama, but considerable research has been conducted within the past 2 decades (e.g., Best and Hudson 1996; Best et al. 1993; Goebel 1996; Henry 1998; Kiser 1996, 2000; Milam 1996; Thomas and Best 2000). Bats occurring on Redstone Arsenal, Madison Co., Alabama, are of special interest because two endangered species are known from the area, M. grisescens and M. sodalis. In preparing management plans, it is desirable to know if either of these species is present and what habitats they may occupy. Goals of this study were to 1) determine species of bats present on Redstone Arsenal, 2) determine habitat associations at 3 spatial scales, 3) elucidate any differences between capture and random sites, 4) and recommend management practices for Redstone Arsenal, especially for endangered species. MATERIALS AND METHODS The study area, Redstone Arsenal, Alabama, includes 15,305 ha in southwestern Madison County. It has been an active military installation since 1941, and it was first used for construction and disposal of chemical-warfare agents. Currently, Redstone Arsenal is headquarters for rocket and missile design and testing by the National Aeronautics and Space Administration (NASA) and the United States Army. Redstone Arsenal is part of the Tennessee Valley physiographic region and is bordered to the south by the Tennessee River. Habitat communities on Redstone Arsenal are upland forest, wet-mesic river floodplain forest, forested palustrine wetlands, springs, and caves (J. C. Godwin and J. L. Hilton, in litt.). I used mist nets to capture bats at 248 sites on Redstone Arsenal in July 2005 (13 sites), May-July 2006 (111 sites), and May?July 2007 (124 4 sites). I attempted to mist net bats in as many areas as possible and selected netting sites in areas where I believed a bat could be captured (Fig. 1.1). Mist nets were placed on or near forest roads, fire breaks, ponds, and creeks. Mist nets ranged in size from 2.6 to 18 m in length, each being 2.6-m high, and size of net used depended upon size of the feature that was netted. I used a double-stacked configuration at most sites where 1 net was stacked on top of another to increase sampling area. For features that were not large enough to contain a double-stacked net, I used a single-high mist net. Where possible, nets were placed so that vegetation surrounded the sides and top of the net to prevent bats from flying around the net. Each net remained open for a total of 6 hours/night (2000- 0200 h). Data on collecting sites, species captured, date and time of capture, sex, age (adult or young-of-year), and reproductive condition, were recorded. Age-group was determined by inspecting the degree of ossification of the epiphyseal plates in joints of the phalanges (Anthony 1988). I collected coordinates for each sampling site using a Global Positioning System (GPSmap76, Garmin International, Inc., Olathe, Kansas). A landscape-level dataset was obtained from the National Land Cover Database (NLCD) 2001 (Limpert et al. 2007); this database describes 21 classes of land-cover types throughout the United States and was created using high-resolution imagery (J. E. Vogelmann et al., http://seamless.usgs.gov/). I merged 13 classes of landcover that occurred on Redstone Arsenal as described by Aebischer et al. (1993) and investigated 8 types of land cover including open water (rivers, streams, lakes, and ponds), developed land (included buildings and roads), deciduous forest, evergreen forest, mixed forest, scrub (present at edge of forest), open pasture, and wetland (marshes and wooded wetlands). I also used 5 an existing ArcMap data layer obtained from the Natural Resources Division at Redstone Arsenal. The UTM coordinates for each sampling site were imported into ArcMap (Environmental Systems Research Institute 2005). I created 250, 500, and 1,000-m buffers i.e., a specified radius around a given point, used to calculate habitat, around each site (Kennedy 2006). These buffers related to average foraging distances of bats captured (Limpert et al. 2007). Buffers were intersected with a layer of land-cover data for the area covered by Redstone Arsenal plus an area 3 km outside the perimeter for buffers that extended beyond the Arsenal. Next, amount of each habitat (in m 2 ) within each buffer was calculated using the calculate-area tool in ArcMap. I also created 250 random points to compare with points where nets were placed; the same procedure for creating buffers and calculating amount of habitat was followed. Area of each habitat in the three buffers was calculated for sites where bats were captured, where bats were not captured, and random points. A species was indicated as present (was captured) or absent (was not captured) at sites where nets were placed. To adequately describe habitats occupied by each species, I performed analyses when ?10 individuals of that species were captured (Aebischer et al. 1993). Habitat associations were tested using multivariate analysis of variance (MANOVA; McCune and Grace 2002). MANOVA was used to detect differences between habitat variables at sites where bats were captured versus where bats were not captured, at sites where bats were captured versus random points, and at sites where an attempt to capture bats was made (includes capture sites and no-capture sites) versus random points. I also performed GLM (General Linear Models) to see which variables were influencing 6 dependent variables. These tests were performed at each spatial scale for each species; alpha-level was P < 0.05. RESULTS During July 2005, May?July 2006, and May?July 2007, 248 sites were sampled and 185 bats were captured, representing 7 species (Table 1.1). Because samples for 6 species were large enough, ecological associations of these species were analyzed statistically. Bats were captured at 82 sites. Although positively identified to species, age, sex, and reproductive condition of 13 individuals were not determined due to their escape from the net. Capture sites versus no-capture sites.--At the 250-m spatial scale, when all capture sites (7 species included) were combined and compared to all no-capture sites, there was a statistically significant difference (Wilk?s ? = 0.934 P = 0.036); developed habitat (F = 7.02, P = 0.009) was more abundant at no-capture sites and wetland habitat (F = 8.91, P = 0.003) was more abundant at capture sites. I also wanted to see if there was any difference between groups, i.e., capture sites and no-capture sites, at the species level, so I performed a MANOVA for each species. I tested for differences between capture and no-capture sites for all sites where a species was captured versus all sites where that species was not captured. All habitat variables differed between capture and no-capture sites for M. austroriparius (Wilk?s ? = 0.926, P = 0.018). Univariate (general linear model; GLM) analysis indicated that the most important differences between capture and no-capture sites were scrub (F = 4.47, P = 0.035) and wetland habitats (F = 11.21, P = 0.001) for M. austroriparius (Table 1.2). When I plotted average area of each habitat type for sites where M. austroriparius was captured and sites where the species 7 was not captured (Fig. 1.3), there was a greater amount of scrub habitat where M. austroriparius was not captured and a greater amount of wetland habitat where the species was captured. When I plotted average areas of each habitat type for all species, I ascertained that, for N. humeralis, there was significantly less developed land (F = 4.42, P = 0.036) and more wetland habitat (F = 6.34, P = 0.013) at sites where N. humeralis was captured resulting in a habitat association with wetland and not with developed land. No significant habitat variable was associated with captures of E. fuscus, L. borealis, M. grisescens, or P. subflavus. At the 500-m spatial scale, when all captures were combined and compared to no- capture sites, there was no statistically significant difference, but developed land (F = 7.02, P = 0.009) was more abundant at no-capture sites. Wilk?s ? was not significant for any species. I plotted average area of each habitat type as done for the 250-m buffer and determined there was a greater amount of evergreen habitat (F = 4.76, P = 0.030) present at sites where L. borealis was captured, indicating a habitat association (Fig. 1.4). Wetland (F = 12.53, P = 0.001) habitat was a significant habitat association for M. austroriparius. Developed land (F = 4.71, P = 0.031) was greater at sites where N. humeralis was not captured, indicating that N. humeralis is not associated with developed land. Open water (F = 4.44, P = 0.036) was associated with capture of P. subflavus and mixed forest (F = 5.06, P = 0.025) habitat was associated with not capturing P. subflavus. There was no significant variable associated with captures of E. fuscus or M. grisescens. At the 1,000-m spatial scale, when all captures were combined and compared to no-capture sites, there was no statistically significant difference, but more open water habitat (F = 5.81, P = 0.017) occurred at capture sites. Habitat variables differed between 8 capture and no-capture sites for one species, M. austroriparius (Wilk?s ? = 0.907, P = 0.003). The univariate portion of this analysis indicated that more open water (F = 4.14, P = 0.043) and wetland (F = 11.67, P = 0.001), and less deciduous forest (F = 4.99, P = 0.026) and mixed forest (F = 5.52, P = 0.020) were statistically significant habitat associations for M. austroriparius (Fig. 1.5). Mixed forest (F = 4.16, P = 0.043) habitat occurred significantly more at sites where P. subflavus was captured, resulting in a habitat association. There was no statistically significant habitat association for E. fuscus, L. borealis, M. grisescens, or N. humeralis. Capture sites versus random sites.--At the 250-m spatial scale, when all captures were combined and compared to random sites, there was a statistically significant difference (Wilk?s ? = 0.891, P < 0.001); deciduous forest (F = 6.73, P = 0.010) was more prevalent at capture sites, while open pasture (F = 4.99, P < 0.001) was more prevalent at random sites. Habitat variables differed between capture and random sites for E. fuscus (Wilk?s ? = 0.935, P = 0.018), L. borealis (Wilk?s ? = 0.932, P = 0.011), and M. austroriparius (Wilk?s ? = 0.935, P = 0.028). The univariate portion of the analysis indicated that presence of more evergreen forest (F = 4.27, P = 0.040), mixed forest (F = 6.16, P = 0.014), and scrub (F = 4.75, P = 0.030) habitats were associated with capture of L. borealis (Fig. 1.6). There was no habitat association for E. fuscus, M. grisescens, P. subflavus, N. humeralis, or M. austroriparius. At the 500-m spatial scale, when all captures were combined and compared to random sites, there was an overall difference (Wilk?s ? = 0.901, P = 0.037), and developed (F = 22.70, P < 0.001), deciduous forest (F = 6.45, P = 0.012), and wetland (F = 17.15, P < 0.001) habitats were significantly different between capture sites and 9 random sites. Overall, there was a greater amount of developed habitat at random sites and a greater amount of deciduous forest and wetland habitats at capture sites. Habitat variables differed between capture and random sites for E. fuscus (Wilk?s ? = 0.943, P = 0.041), L. borealis (Wilk?s ? = 0.933, P = 0.035), P. subflavus (Wilk?s ? = 0.938, P = 0.034), and M. austroriparius (Wilk?s ? = 0.907, P = 0.01). The univariate portion of analyses indicated that developed land occurred more at random sites for E. fuscus (F = 8.35, P < 0.001), L. borealis (F = 10.28, P = 0.041), P. subflavus (F = 6.34, P = 0.012), N. humeralis (F = 8.36, P = 0.004), and M. austroriparius (F = 5.68, P = 0.018). Wetland habitat was associated with capture of E. fuscus (F = 6.91, P = 0.009), P. subflavus (F = 7.22, P = 0.008), N. humeralis (F = 8.73, P = 0.003), and M. austroriparius (F = 23.27, P < 0.001). Deciduous forest (F = 5.13, P = 0.024) was significantly associated with captures of L. borealis. There was no habitat association related to captures of M. grisescens. At the 1,000-m spatial scale, when all captures were combined and compared to random sites, there was an overall difference (Wilk?s ? = 0.9116, P < 0.001), and open water (F = 4.37, P = 0.037), deciduous forest (F = 4.92, P = 0.027), and wetland (F = 15.54, P < 0.001) were more prevalent at capture sites rather than random sites. Developed (F = 14.31, P < 0.001) and open pasture (F = 5.48, P = 0.020) habitats were more abundant at random sites. Habitat variables differed between capture and random sites for E. fuscus (Wilk?s ? = 0.889, P < 0.001), L. borealis (Wilk?s ? = 0.939, P = 0.024), P. subflavus (Wilk?s ? = 0.932, P = 0.020), N. humeralis (Wilk?s ? = 0.939, P = 0.036), and M. austroriparius (Wilk?s ? = 0.889, P < 0.001). Univariate analyses indicated that area open water was greater at sites where E. fuscus (F = 4.10, P = 0.044) 10 and M. austroriparius (F = 4.10, P = 0.044) were captured versus random sites, indicating that these species are associated with open water. Developed land was significantly greater at random sites for E. fuscus (F = 4.41, P = 0.037), L. borealis (F = 6.57, P = 0.011), P. subflavus (F = 4.27, P = 0.040), N. humeralis (F = 6.57, P = 0.011), and M. austroriparius (F = 4.41, P = 0.034), resulting in no habitat association between these species and developed land. Wetland habitat was associated with E. fuscus (F = 24.44, P < 0.001), M. grisescens (F = 4.09, P = 0.044), P. subflavus (F = 10.72, P = 0.001), N. humeralis (F = 8.02, P = 0.005), and M. austroriparius (F = 24.44, P < 0.001). Deciduous forest was significantly associated with captures of L. borealis (F = 4.58, P = 0.033). Capture and no-capture sites versus random sites.--Habitat variables differed between capture plus no-capture sites and random sites at the 250-m (Wilk?s ? = 0.880, P < 0.001), 500-m (Wilk?s ? = 0.896, P < 0.001), and 1,000-m (Wilk?s ? = 0.921, P < 0.001) spatial scales. Univariate analyses indicated that deciduous forest (F = 17.38, P < 0.001), mixed forest (F = 7.94, P = 0.005), and scrub (F = 8.38, P = 0.004) occurred more often at net sites and open pasture (F = 9.30, P = 0.002) occurred more often at random sites at the 250-m scale. Area of deciduous forest (F = 12.84, P < 0.001) and wetland (F = 15.06, P < 0.001) occurred in greater amounts at net sites versus random sites while area of developed (F = 31.81, P < 0.001) and open pasture (F = 4.41, P = 0.036) occurred in greater amount at random sites at the 500-m scale. Presence of deciduous forest (F = 12.46, P < 0.001), scrub (F = 4.00, P = 0.046), and wetland (F = 12.33, P < 0.001) habitat were significantly greater at net sites, and developed (F = 20.09, 11 P < 0.001) and open pasture (F = 5.64, P = 0.018) habitats were significantly greater at random sites at the 1,000-m-scale. DISCUSSION Comparison of capture versus no-capture sites demonstrated that I caught more bats in areas where open water and wetland habitats occurred more frequently and significantly fewer at sites primarily containing developed land. A study by Sparks et al. (2005) determined that bats avoided developed land while foraging. This could be because there are fewer insects and less diversity of insects in developed areas as opposed to rural areas (Geggie and Fenton 1985; Sparks et al. 2005). Duchamp et al. (2004) also reported that bats (E. fuscus and N. humeralis) selected more wooded habitats than developed habitats as foraging areas. Contrary to my study, they reported that these species avoided open water when foraging, but Duchamp et al. (2004) did not define creeks and streams as open water while I did. Based on comparison of capture versus random sites, significantly more bats were captured in areas that had a greater amount of deciduous forest and wetland habitat and a lesser amount of developed land and open pasture. Bats frequently use deciduous forest (Barbour and Davis 1969; Davis and Mumford 1962; Fujita and Kunz 1984; LaVal et al. 1977) for foraging and less frequently use wetland habitat (Barbour and Davis 1969) for foraging. L. borealis (Shump and Shump 1982), E. fuscus, and N. humeralis (Duchamp et al. 2004) often forage in open fields. Comparison of capture and no-capture sites versus random sites verified that I did not sample randomly and that I chose to set nets at sites that had more deciduous forest, mixed forest, scrub, and wetland habitats, while random sites had more developed land 12 and open pasture. It is possible that habitat associations I detected were influenced by placement of nets. A way to remedy this in future studies would be to randomly place nets; however, when surveying for bats, success of capture decreases with random placement of nets, because it is difficult to catch bats in nets that are not conducive to capture, i.e., nets placed in a pasture or in the middle of a cluttered forest lack the structural component of canopy that funnels bats into the net. Bats are more easily captured in nets placed across flyways or over water (Kunz and Kurta 1988). Sites that are conducive to capturing bats include those that are at a water source or across paths used as flyways (Kunz and Kurta 1988). Because I did not randomly place nets, I have more confidence in habitat associations I ascertained using comparison of capture versus no-capture sites as compared to comparison of capture versus random sites. Absence of bats at a site may be influenced by factors other than habitat surrounding the site. For example, fog reduces foraging activity by bats (Pye 1971). Phase of the moon may (Fenton et al. 1977) or may not (Hayes 1997) influence activity of bats. Also, as light intensity of the moon increases, bats forage higher in the canopy (Fenton et al. 1977; Hecker and Brigham 1999). This does not directly affect whether or not bats forage but where they forage. During my study, it was possible that on moonlit nights bats may have moved higher in the canopy above my mist nets. A more probable cause for absence of bats at a site is a decrease in abundance of insects. Activity of bats decreases as biomass of insects decreases; therefore, if insects are not available, activity of bats will be limited to drinking and commuting (Hayes 1997). Redstone Arsenal was used as foraging habitat by 1 of the 2 endangered species of bats that occur in Alabama. No M. sodalis was captured, but 11 M. grisescens of both 13 sexes were captured on the Arsenal. One female was post-lactating (Table 1.1), which indicated M. grisescens was reproducing on or near Redstone Arsenal. M. grisescens born in summer 2006 also were captured in 2006. It is likely that M. grisescens roosts outside of Redstone Arsenal near the Tennessee River (Tuttle 1976), although it is possible that M. grisescens roosts on Redstone Arsenal. Clearly, Redstone Arsenal is within the foraging range of this endangered species. Most M. grisescens captured were at sites where mist nets were over water, but 4 were captured over roadways surrounded by deciduous forest that acted as flyways to riparian areas. M. grisescens forages over rivers and lakes, and probably uses small creeks and streams as foraging areas or as flyways to get to larger bodies of water to forage (Best and Hudson 1996; Harvey et al. 1999). My study was unable to detect a specific habitat association for M. grisescens, but this species forages in riparian areas (LaVal et al. 1977) and uses forest canopy as protection against predation (Tuttle 1976). Number of captures increased toward the end of summer (Fig. 1.2). This probably is related to the time that young become volant. Young bats are able to fly 3-4 weeks after they are born (Decher and Choate 1995; Best 2004a), and as inexperienced flyers, they are relatively easy to catch in mist nets. My study did not detect a significant habitat association for E. fuscus, but previous studies have shown E. fuscus to be a habitat generalist (Agosta 2002; Kurta and Baker 1990). L. borealis is associated with evergreen forest and is known to fly above tree canopy and over open pastures (LaVal et al. 1977; Shump and Shump 1982). I suspect that L. borealis is also a habitat generalist when foraging because I captured the species in many habitats. M. austroriparius foraged in open water and wetland habitats, 14 similar to habitats described by Barbour and Davis (1969). M. austroriparius was associated with areas that had less scrub, deciduous forest, and mixed forest. N. humeralis was associated with wetland habitats, and away from developed land. The species forages in woodlands and roosts in buildings (Watkins 1972), but has not been documented to forage in wetlands. I frequently captured N. humeralis foraging over ponds. P. subflavus foraged over open water and it was associated with areas containing less mixed forest. This species uses streams and forest edges as foraging habitat (Davis and Mumford 1962; Fujita and Kunz 1984; LaVal et al. 1977). Species of bats that were not captured on Redstone Arsenal, but whose geographic ranges include the Arsenal are listed in Table 1.4. More sampling sites or different sampling techniques, such as ultrasonic-detection systems, could be employed to increase chances of detecting these species. There are 4 species of bats that my study did not detect that were recorded outside of my study area in Jackson County, a county adjacent to Madison County. I observed Corynorhinus rafinesquii roosting in a cave, and Lasionycteris noctivagans, Lasiurus cinereus, and Myotis sodalis were captured in mist nets in autumn 2008 (pers. observ.). If sampling at Redstone Arsenal extended into autumn, it is possible that these species would be detected. There is no recent record of M. lucifugus from Alabama, and more research on this species is needed (Best 2004c). Overall, this study provided data for use by land managers. This study may also provide a time-effective and cost-effective method for collecting and analyzing habitat data by using GIS techniques rather than collecting vegetation data in the field. MANOVA tests indicated that sites where I placed nets were statistically different from random sites at 3 15 spatial scales. Sites I chose had greater amounts of open water, deciduous forest, and wetland habitats, which probably are more desirable habitats for bats. Management Implications.--One endangered species, M. grisescens, was captured on Redstone Arsenal during this study. It also was captured on the Arsenal in the past (J. C. Godwin and J. L. Hilton, in litt.). Because M. grisescens is associated with wetlands, and because other studies have ascertained that the species is associated with rivers and lakes (Best and Hudson 1996) and riparian areas in general (LaVal et al. 1977), I suggest that these areas be preserved for foraging habitat for M. grisescens. I recommend that riparian areas on Redstone Arsenal be preserved as foraging habitat for all bats and that amount of lands to be developed be kept to a minimum. Future studies are needed to investigate microhabitat variables important to bats on Redstone Arsenal. It also is necessary to have information about habitats bats are using year-round as opposed to just one season of monitoring (Ball 2002). LITERATURE CITED AEBISCHER, N. J., P. A. ROBERTSON, AND R. E. KENWARD. 1993. Compositional analysis of habitat use from animal radio-tracking data. Ecology 74:1313-1325. AGOSTA, S. J. 2002. Habitat use, diet and roost selection by the big brown bat (Eptesicus fuscus) in North America: a case for conserving an abundant species. Mammal Review 32:179-198. ANTHONY, E. L. P. 1988. Age determination in bats. Pp. 47-58 in Ecological and behavioral methods for the study of bats (T. H. Kunz, ed.). Smithsonian Institution Press, Washington, D.C. BALL, L. C. 2002. A strategy for describing and monitoring bat habitat. Journal of 16 Wildlife Management 66:1148-1153. BARBOUR, R. W., AND W. H. DAVIS. 1969. Bats of America. University Press of Kentucky, Lexington. BEST, T. L. 2004a. Mammals. Pp. 167-169, in Alabama wildlife: imperiled amphibians, reptiles, birds, and mammals (R. E. Mirarchi, M. A. Bailey, T. M. Haggerty, and T. L. Best, eds.). University of Alabama Press, Tuscaloosa, 3:1-225. BEST, T. L. 2004b. Mammals. Pp. 185-204, in Alabama wildlife: a checklist of vertebrates and selected invertebrates: aquatic mollusks, fishes, amphibians, reptiles, birds, and mammals (R. E. Mirarchi, ed.). University of Alabama Press, Tuscaloosa, 1:1-209. BEST, T. L. 2004c. Little brown myotis Myotis lucifugus (LeConte). Pp. 193-194, in Alabama wildlife: imperiled amphibians, reptiles, birds, and mammals (R. E. Mirarchi, M. A. Bailey, T. M. Haggerty, and T. L. Best, eds.). University of Alabama Press, Tuscaloosa, 3:1-225. BEST, T. L., AND M. K. HUDSON. 1996. Assessment of routes used by female gray bats (Myotis grisescens) between roost sites and foraging areas in northern Alabama. Journal of the Alabama Academy of Science 67:6-14. BEST, T. L., S. D. CAREY, K. G. CAESAR, AND T. H. HENRY. 1993. Distribution and abundance of bats (Mammalia: Chiroptera) in the Coastal Plain caves of southern Alabama. National Speleological Society Bulletin 54:61-65. BRIGHAM, R. M. 2007. Bats in forests: what we know and what we need to learn, Pp. 1-15, in Bats in forests: conservation and management (M. J. Lacki, J. P. 17 Hayes, and A. Kurta, eds.). Johns Hopkins University Press, Baltimore, Maryland. DAVIS, W. H., AND R. E. MUMFORD. 1962. Ecological notes on the bat Pipistrellus subflavus. American Midland Naturalist 68:394-398. DECHER, J., AND J. R. CHOATE. 1995. Myotis grisescens. Mammalian Species 510:1-7. DUCHAMP, J. E., D. W. SPARKS, AND J. O. WHITAKER. 2004. Foraging-habitat selection by bats at an urban-rural interface: comparison between a successful and a less successful species. Canadian Journal of Zoology 82:1157-1164. ENVIRONMENTAL SYSTEMS RESEARCH INSTITUTE. 2005. ArcGIS desktop and workstation software, version 9.1. Environmental Systems Research Institute, Redlands, California. EVELYN, M. J., AND D. A. STILES. 2007. Roosting requirements of two frugivorous bats (Sturnira lilium and Arbiteus intermedius) in fragmented Neotropical forest. Biotropica 35:405-418. FAHRIG, L. 2003. Effects of habitat fragmentation on biodiversity. Annual Review of Ecology, Evolution, and Systematics 34:487-515. FENTON, M. B., N. G. H. BOYLE, T. M. HARRISON, AND D. J. OXLEY. 1977. Activity patterns, habitat use, and prey selection by some African insectivorous bats. Biotropica 9:73-85. FUJITA, M. S., AND T. H. KUNZ. 1984. Pipistrellus subflavus. Mammalian Species 228:1-6. 18 GEGGIE, J. F., AND M. B. FENTON. 1985. A comparison of foraging by Eptesicus fuscus (Chiroptera: Vespertilionidae) in urban and rural environments. Canadian Journal of Zoology 63:263-267. GOEBEL, A. B. 1996. Temporal variation in movement patterns of adult female Myotis grisescens (Chiroptera: Vespertilionidae). M.S. thesis, Auburn University, Alabama. GORRESEN, P. M., AND M. R. WILLIG. 2004. Landscape responses of bats to habitat fragmentation in Atlantic Forest of Paraguay. Journal of Mammalogy 85:688- 697. GORRESEN, P. M., M. R. WILLIG, AND R. E. STRAUSS. 2005. Multivariate analysis of scale-dependent associations between bats and landscape structure. Ecological Applications 15:2126-2136. HALL, E. R. 1981. The mammals of North America. John Wiley and Sons, New York. HARVEY, M. J., J. S. ALTENBACH, AND T. L. BEST. 1999. Bats of the United States. Arkansas Game and Fish Commission and United States Fish and Wildlife Service, Little Rock, Arkansas. HAYES, J. P. 1997. Temporal variation in activity of bats and the design of echolocation monitoring studies. Journal of Mammalogy 78:514-524. HECKER, K. R., AND R. M. BRIGHAM. 1999. Does moonlight change vertical stratification of activity by forest-dwelling insectivorous bats? Journal of Mammalogy 80:1196-1201. HENRY, T. H. 1998. Variation in use of habitats by the gray bat (Myotis grisescens) in northern Alabama. M.S. thesis, Auburn University, Alabama. 19 KALCOUNIS-R?PPELL, M. C., J. M. PSYLLAKIS, AND R. M. BRIGHAM. 2005. Tree roost selection by bats: an empirical synthesis using meta-analysis. Wildlife Society Bulletin 33:1123-1132. KALCOUNIS, M. C., K. A. HOBSON, R. M. BRIGHAM, AND K. R. HECKER. 1999. Bat activity in the boreal forest: importance of stand type and vertical strata. Journal of Mammalogy 80:673-682. KENNEDY, M. 2006. Introducing geographic information systems with ArcGIS. John Wiley and Sons, Inc., Hoboken, New Jersey. KISER, W. M. 1996. Conservation of LeConte's free-tailed bat (Tadarida brasiliensis cynocephala): environmental parameters of a natural and an artificial roost. M.S. thesis, Auburn University, Alabama. KISER, W. M. 2000. Distribution and status of LeConte?s free-tailed bat (Tadarida brasiliensis cynocephala) in Alabama. Occasional Papers of the North Carolina Museum of Natural Sciences and the North Carolina Biological Survey 12:67-73. KUNZ, T. H., AND A. KURTA. 1988. Capture methods and holding devices. Pp. 1-29 in Ecological and behavioral methods for the study of bats (T. H. Kunz, ed.). Smithsonian Institution Press, Washington, D. C. KURTA, A., AND R. H. BAKER. 1990. Eptesicus fuscus. Mammalian Species 356:1-10. LAVAL, R. K., R. L. CLAWSON, M. L. LAVAL, AND W. CAIRE. 1977. Foraging behavior and nocturnal activity patterns of Missouri bats, with emphasis on the endangered species Myotis grisescens and Myotis sodalis. Journal of Mammalogy 58:592- 599. 20 LIMPERT, D. L., D. L. BIRCH, M. S. SCOTT, M. ANDRE, AND E. GILLAM. 2007. Tree selection and landscape analysis of eastern red bat day roosts. Journal of Wildlife Management 71:478-486. MCCUNE, B., AND J. B. GRACE. 2002. Analysis of ecological communities. MjM Software Design, Gleneden Beach, Oregon. MILAM, B. A. 1996. Daily and seasonal ranges of temperatures of a roost used by the Brazilian free-tailed bat (Tadarida brasiliensis) and the big brown bat (Eptesicus fuscus) in Alabama. M.S. thesis, Auburn University, Alabama. MILES, A. C., S. B. CASTLEBERRY, D. A. MILLER, AND L. M. CONNER. 2006. Multi- scale roost-site selection by evening bats on pine-dominated landscapes in Southwest Georgia. Journal of Wildlife Management 70:1191-1199. MILLER, D. A., E. B. ARNETT, AND M. J. LACKI. 2003. Habitat management for forest- roosting bats of North America: a critical review of habitat studies. Wildlife Society Bulletin 31:30-44. PATRIQUIN, K. J., AND R. M. R. BARCLAY. 2003. Foraging by bats in cleared, thinned and unharvested boreal forest. Journal of Applied Ecology 40:646-657. PYE, J. D. 1971. Bats and fog. Nature 229:572-574. SHUMP, K. A., JR., AND A. U. SHUMP. 1982. Lasiurus borealis. Mammalian Species 183:1-6. SPARKS, D. W., C. M. RITZI, J. E. DUCHAMP, AND J. O. WHITAKER. 2005. Foraging habitat of the Indiana bat (Myotis sodalis) at an urban-rural interface. Journal of Mammalogy 86:713-718. 21 THOMAS, D. P., AND T. L. BEST. 2000. Radiotelemetric assessment of movement patterns of the gray bat (Myotis grisescens) at Guntersville Reservoir, Alabama. Occasional Papers of the North Carolina Museum of Natural Sciences and the North Carolina Biological Survey 12:27-39. TUTTLE, M. D. 1976. Population ecology of the gray bat (Myotis grisescens): factors influencing growth and survival of newly volant young. Ecology 57:587-595. WATKINS, L. C. 1972. Nycticeius humeralis. Mammalian Species 23:1-4. 22 ____________________________________________________________________________________________________________ Table 1.1. Species, sample size (n), gender, age-class, and reproductive status of bats captured at Redstone Arsenal, Madison Co., Alabama: YOY, young-of-the-year; NN, not pregnant and not lactating; L, lactating; PL, post-lactating; PR, pregnant; I, immature; S, scrotal; NS, non-scrotal; U, undetermined. ____________________________________________________________________________________________________________ Species n Gender Age Reproductive status Male Female Adult YOY NN L PL PR I S NS U Eptesicus fuscus 53 13 39 42 8 11 16 6 1 4 8 5 1 Lasiurus borealis 54 16 28 23 21 1 5 9 1 12 12 4 10 Myotis austroriparius 15 6 9 9 6 - 5 2 - 2 1 5 - Myotis grisescens 11 9 2 7 4 - - 1 - 1 2 7 - Myotis septentrionalis 2 2 - 1 1 - - - - - - 2 - Nycticeius humeralis 33 17 15 28 4 4 4 7 - 1 15 2 1 Perimyotis subflavus 16 11 5 12 3 1 2 1 - 1 2 9 - Unidentified Myotis 1 - - - - - - - - - - - 1 Total 185 ____________________________________________________________________________________________________________ 23 __________________ Table 1.2. Significant habitat associations of bats on Redstone Arsenal, Madison Co., Alabama, 2005-2007, as determined by GLM comparing mean area of each habitat type where a species was captured versus sites where it was not captured. A positive sign indicates habitats where bats were captured and a negative sign indicates habitat where bats were not captured. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. ____________________________________________________________________________________________________________ Variable and Size of Buffer Species Lasiurus borealis Evergreen 500-m (+)* Myotis austroriparius Shrub/Scrub 250-m (-)* Wetland 250-m (+)** Wetland 500-m (+)** Open Water 1,000-m (+)* Deciduous 1,000-m (-)* Mixed Forest 1,000-m (-)* Wetland 1,000-m (+)** Nycticeius humeralis Developed 250-m (-)* Wetland 250-m (+)* Developed 500-m (-)* Perimyotis subflavus Open Water 500-m (+)* Mixed Forest 500-m (-)* Mixed 1,000-m (-)* __________________________________________________________________________________________________ __________________________________________________________________________________________________________ Table 1.3. Significant habitat associations for bats on Redstone Arsenal, Madison Co., Alabama, 2005-2007, as determined by GLM comparing mean area of each habitat type at sites where a species was captured against random sites. A positive sign indicates habitats where bats were detected and a negative sign indicates habitats where bats were not detected. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. ____________________________________________________________________________________________________________ ____________________________________________________________________________________________________________ Variable and Size of Buffer Species Eptesicus fuscus Developed 500-m (-)** Wetland 500-m (+)* Open Water 1,000-m (+)* Developed 1,000-m (-)* Wetland 1,000-m (+)** Lasiurus borealis Evergreen Forest 250-m (+)* Mixed Forest 250-m (+)* Shrub/Scrub 250-m (+)* Developed 500-m (-)** Deciduous Forest 500-m (+)* Developed 1,000-m (-)** Deciduous Forest 1,000-m (+)* Myotis austroriparius Developed 500-m (-)* Wetland 500-m (+)** Open Water 1,000-m (+)* Developed 1,000-m (-)* Wetland 1,000-m (+)** Nycticeius humeralis Developed 500-m (-)** Wetland 500-m (+)** Developed 1,000-m (-)* Wetland 1,000-m (+)* Perimyotis subflavus Developed 500-m (-)* Wetland 500-m (+)* Developed 1,000-m (-)* Wetland 1,000-m (+)** 24 25 Table 1.4. Species of bats that were not captured at Redstone Arsenal, Madison Co., Alabama, but potentially occur there. ___________________________________________________________________________________________________ Corynorhinus rafinesquii Lasionycteris noctivagans Lasiurus cinereus Lasiurus seminolus Myotis lucifugus Myotis sodalis Tadarida brasiliensis ___________________________________________________________________________________________________ Fig. 1.1 Sites where mist nets were placed to capture bats on Redstone Arsenal, Madison, Co., Alabama, summers 2005-2007. 26 Fig. 1.2 Dates bats were captured at Redstone Arsenal, Madison Co., Alabama. These data indicate that number of bats captured increased as summers progressed during 2005- 2007. Captures for 2005-2007 0 5 10 15 20 25 30 35 40 Ma y W e e k 2 M a y W e e k 3 M a y W ee k 4 Ju n e W e e k 1 J u n e We e k 2 Ju n e W e e k 3 J u n e We e k 4 J u ly W e e k 2 Ju l y W e e k 3 J u ly W e e k 4 Sampling Period Nu m b e r o f B a ts 27 Fig. 1.3 Average area (m 2 ) of habitat types at the 250-m spatial scale at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates capture sites and a striped bar indicates no-capture sites for a) Eptesicus fuscus, b) Lasiurus borealis, c) Myotis austroriparius, d) M. grisescens, e) Nycticeius humeralis, and f) Perimyotis subflavus. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. 28 Fig. 1.4 Average area (m 2 ) of habitat types at the 500-m spatial scale at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates capture sites and a striped bar indicates no-capture sites for a) Eptesicus fuscus, b) Lasiurus borealis, c) Myotis austroriparius, d) M. grisescens, e) Nycticeius humeralis, and f) Perimyotis subflavus. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. 29 Fig. 1.5 Average area (m 2 ) of habitat types at the 1,000-m spatial scale at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates capture sites and a striped bar indicates no-capture sites for a) Eptesicus fuscus, b) Lasiurus borealis, c) Myotis austroriparius, d) M. grisescens, e) Nycticeius humeralis, and f) Perimyotis subflavus. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. 30 Fig. 1.6 Average area (m 2 ) of habitat types at the 250-m spatial scale at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates capture sites and a striped bar indicates random sites for a) Eptesicus fuscus, b) Lasiurus borealis, c) Myotis austroriparius, d) M. grisescens, e) Nycticeius humeralis, and f) Perimyotis subflavus. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. 31 Fig. 1.7 Average area (m 2 ) of habitat types at the 500-m spatial scale at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates capture sites and a striped bar indicates random sites for a) Eptesicus fuscus, b) Lasiurus borealis, c) Myotis austroriparius, d) M. grisescens, e) Nycticeius humeralis, and f) Perimyotis subflavus. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. 32 Fig. 1.8 Average area (m 2 ) of habitat types at the 1,000-m spatial scale at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates capture sites and a striped bar indicates random sites for a) Eptesicus fuscus, b) Lasiurus borealis, c) Myotis austroriparius, d) M. grisescens, e) Nycticeius humeralis, and f) Perimyotis subflavus. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. 33 Fig. 1.9 Average area (m 2 ) of habitat types at the a) 250-m, b) 500-m, and c) 1,000-m spatial scale at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates sites where nets were placed (captures and no-captures) and a striped bar indicates random sites. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. 34 35 CHAPTER 2: HABITAT ASSOCIATIONS AMONG BATS ON REDSTONE ARSENAL, MADISON CO., ALABAMA, AS DETERMINED BY ULTRASONIC-DETECTION SURVEYS ABSTRACT Ultrasonic detection of bats may allow the observer to detect species that are less frequently captured in mist nets. In this study, I used AnaBat SD1 detectors to obtain species identification at locations in northern Alabama. I used multivariate analysis to evaluate associations between habitat types and species using those habitats at 3 spatial scales. Results obtained from this study suggested the following associations: the eastern red bat (Lasiurus borealis) with open water, gray myotis (Myotis grisescens) with open water and deciduous forest, evening bat (Nycticeius humeralis) with open water, and perimyotis (Perimyotis subflavus) with open water and wetlands. INTRODUCTION Acoustic-monitoring devices may be more reliable in identifying bats in an area of interest because there is a potential bias in using mist nets for studies of foraging ecology. Some species are more easily captured by mist nets, while fast-flying, high- altitude species may not be captured in mist nets (Lacki et al. 2007). For example, the Brazilian free-tailed bat (Tadarida brasiliensis) is a high-flying species that occurs in Alabama (Wilkins 1989). This species, as well as others, are detectable ultrasonically 36 and have been monitored using acoustic-monitoring devices in the southeastern United States (Britzke 2003). AnaBat (Titley Electronics, Ballina, New South Wales, Australia) detectors use frequency-division to make echolocation calls by bats audible to humans. The Zero- Crossing Analysis Interface Module (ZCAIM: Titley Electronics, Ballina, New South Wales, Australia) of AnaBat records the call and allows the observer to graphically observe calls (Fenton et al. 2001). Call files are downloaded to a computer from a compact flash (CF) card and can be analyzed using the DOS-operated program Analook (Version 4.9 j: Titley Electronics, Ballina, New South Wales, Australia). It is possible to analyze variables such as frequency, wavelength, duration, and slope. Measures of these parameters are unique for each species of bat, so that comparisons can be made among calls that were collected in the field and calls of known species of bats (Britzke 2003). There are 2 methods of monitoring echolocation calls of bats; actively and passively. Active monitoring is done when the observer uses an acoustic-monitoring device to collect calls by directing the device toward a bat that is flying overhead. Active monitoring allows the observer to collect long, high-quality calls and identify bats on the wing but requires an observer to be present with the acoustic-monitoring device at all times. For passive monitoring, acoustic-monitoring devices are left overnight or for multiple nights, usually in a weather-proof container, and parameters of calls are stored for later analysis (Murray et al. 1999). Passive monitoring is used more often for studies of use of habitats because multiple acoustic-monitoring devices can collect data over a longer period than with active monitoring. Several studies have used AnaBat, an ultrasonic-detection system, as a means of investigating habitat use by bats (Erickson and 37 West 2003; Glendell and Vaughan 2002; Kalcounis et al. 1999; Loeb and O'Keefe 2006; Vaughan et al. 1997; Yates and Muzika 2006). In this study, I used AnaBat to assess habitat relationships among a community of bats in northern Alabama. MATERIALS AND METHODS The study area was Redstone Arsenal, Madison Co., Alabama, which was described in Chapter 1. I used 3 AnaBat SD1 ultrasonic detectors (Titley Electronics, Ballina, New South Wales, Australia) to detect bats at 93 sites during May-July 2007. All detectors were deployed 3nights/week at different sites, and each was moved to a new location the following night. I attempted to place detectors in as many areas of Redstone Arsenal as possible and selected sites in areas where I believed a bat could be detected (Fig. 2.1). Features where detectors were placed included fly-ways along roads and creeks, open fields, and ponds. Detectors were contained in a weatherproof housing and directed toward a Lexan reflector placed at a 45? angle to limit interference of rain with the detector?s microphone (Fig. 2.2; modified from Yates and Muzika 2006). Detectors were placed ca. 1.5 m from the ground on a tree, fence-post, or utility pole. Each detector was set at a sensitivity of 7.5 and programmed to record from 1800 to 0600 h CDT. Files were analyzed using discriminant-function analysis as described by Britzke (2003). Data from detectors were downloaded to a personal computer and viewed using Analook version 4.9j. I used a filter created by E. Britzke (pers. comm.) to remove ultrasonic calls that had <5 pulses in a sequence. This filter also removed files created by noise of wind and insects. Parameters of calls were imported into MINITAB 14, and I performed discriminant-function analysis. For a species to be considered present at a site, there had to be ?2 sequences from a species (because pulses from one species of bat 38 can resemble the call of another species), where ?50% of pulses in the sequence were identified as that species (modified from Samoray 2002). I defined a valid call as one that had ?5 pulses in a sequence, where there were ?2 sequences from a species at each site with ?50% of pulses in the sequence identified as that species (modified from Samoray 2002). The same procedure for importing data, creating buffers, and calculating habitat was performed as outlined in Chapter 1. I also used 250 random points generated as described in Chapter 1 to make comparisons with points where bats were detected and for creating habitat analyses. Area of each habitat in the 3 buffers was calculated for sites where bats were detected, where bats were not detected, and at random points. A species was indicated as either present (was detected) or absent (was not detected). I performed analyses for species only when calls of that species were detected at ?10 sites (Aebischer et al. 1993). Habitat associations were tested using multivariate analysis of variance (MANOVA; McCune and Grace 2002). MANOVA was used to determine if there was a difference between habitat variables at sites where bats were detected versus where bats were not detected, at sites where bats were detected versus random points, and at sites where an attempt to detect bats was made (includes detection sites and no-detection sites) versus random points. These tests were performed at each spatial scale for each species; alpha-level was P < 0.05. RESULTS During May?July 2007, I recorded calls at 93 sites. Over the sampling season, 104 valid identifications of bats were ascertained at 45 of the sites; these identified calls represented 6 species. Species identified were the big brown bat (Eptesicus fuscus), 39 eastern red bat (Lasiurus borealis), gray myotis (Myotis grisescens), northern myotis (M. septentrionalis), evening bat (Nycticeius humeralis), and perimyotis (Perimyotis subflavus). Five of the species had a sample size ?10 and were used in statistical analyses (M. septentrionalis was excluded). Detection sites versus no-detection sites.--At the 250-m spatial scale, when detections of all species were combined and compared to no-detection sites, there was an overall difference (Wilk?s ? = 0.811, P = 0.020). Open water (F = 8.52, P = 0.004) and deciduous forest (F = 5.14, P = 0.026) occurred in greater amounts where bats were detected, and developed land (F = 4.37, P = 0.039) occurred more at sites where bats were not detected. Habitat variables differed between detection and no-detection sites for two species, M. grisescens (Wilk?s ? = 0.605, P < 0.001) and P. subflavus (Wilk?s ? = 0.633, P < 0.001). This indicates that overall, habitat at detection sites was statistically different from habitat at no-detection sites for these two species. The univariate (general linear model; GLM) portion of the analysis indicated that area of open water (F = 8.42, P = 0.005) was significantly different between sites where L. borealis was detected versus sites where L. borealis was not detected. When I plotted average area of open water at detection sites and at no-detection sites, I determined that a greater area open water occurred at sites where L. borealis was captured. I plotted average area for all types of habitat at detection versus no-detection sites to make habitat associations for each species. Open water (F = 17.48, P < 0.001) and deciduous forest (F = 13.75, P < 0.001) were associated with detection of M. grisescens (Table 2.2). Developed land (F = 5.25, P = 0.024) and open pasture (F = 12.76, P < 0.001) were associated with sites where M. grisescens was not detected. For N. humeralis, open water (F = 7.90, P = 0.006) was a 40 habitat association. Open water (F = 21.38, P < 0.001) and wetland (F = 11.46, P = 0.001) were associated with detection of P. subflavus. Developed (F = 10.14, P = 0.002), mixed forest (F = 4.79, P = 0.031), and open pasture (F = 7.01, P = 0.010) habitats were associated with sites where P. subflavus was not detected. There was no significant habitat association for E. fuscus. At the 500-m-spatial scale, when habitat variables from all sites where bats were detected were combined and compared to no-detection sites, there was no overall difference, but open water (F = 5.83, P = 0.018) and deciduous forest (F = 4.68, P = 0.033) occurred more at detection sites while developed habitat (F = 4.90, P = 0.029) occurred more at no-detection sites. Habitat variables differed between detection and no- detection sites for two species, M. grisescens (Wilk?s ? = 0.679, P < 0.001) and P. subflavus (Wilk?s ? = 0.768, P = 0.003). The univariate portion of this analysis indicated that open water (F = 9.96, P = 0.002) was significantly greater for sites where L. borealis was detected. Open water (F = 11.80, P = 0.001), developed (F = 6.57, P = 0.012), deciduous forest (F = 19.08, P < 0.001), and open pasture (F = 10.19, P = 0.001) were significantly different between detection and no-detection sites for M. grisescens. Open water and deciduous forest occurred more where M. grisescens was,detected, and developed land and open pasture occured more where M. grisescens was not detected. Significant habitat associations for P. subflavus were open water (F = 9.59, P = 0.003) and wetland (F = 8.98, P = 0.004). Developed lands (F = 11.40, P = 0.001) and mixed forest (F = 4.44, P = 0.038) were less abundant where the species was detected. There was no significant habitat association for E. fuscus or N. humeralis. 41 At the 1,000-m spatial scale, when all detections were combined and compared to no-detection sites, there was no statistically significant difference, and no habitat variable was significant. Habitat variables differed between detection and no-detection sites for M. grisescens (Wilk?s ? = 0.703, P < 0.001) and P. subflavus (Wilk?s ? = 0.788, P = 0.008). The univariate portion of this analysis and plots of average amounts of type of habitat for detection sites and no-detection sites indicated that deciduous forest (F = 23.38, P < 0.001) was associated with detection of M. grisescens. Developed land (F = 4.83, P = 0.031) and open pasture (F = 8.96, P = 0.004) habitats occurred more where M. grisescens was not detected and were not habitat associations for M. grisescens. Open water (F = 6.99, P = 0.010) and wetland (F = 6.97, P = 0.010) were habitat associations for P. subflavus. Developed lands (F = 10.41, P = 0.002) and mixed forest (F = 4.34, P = 0.040) occurred more in no-detection sites and were not associated with capture of this species. There was no significant habitat association for E. fuscus, L. borealis, or N. humeralis. Detection sites versus random sites.--When all detections were combined and compared to random sites at the 250-m spatial scale, there was no overall statistical difference and no single habitat variable was significant. Habitat variables differed overall between detection and random sites for M. grisescens (Wilk?s ? = 0.935, P = 0.021) and P. subflavus (Wilk?s ? = 0.937, P = 0.022). The univariate portion of the analysis and plots of average amounts of habitat types indicated that presence of open water was associated with detections of M. grisescens (F = 7.70, P = 0.006), N. humeralis (F = 4.61, P = 0.033), and P. subflavus (F = 8.11, P = 0.005). Deciduous forest was associated with detection of M. grisescens (F = 8.20, P = 0.005), while amount of open 42 pasture (F = 6.12, P = 0.014) was greater at random sites and was not a habitat association. There was no statistically significant habitat variable for E. fuscus or L. borealis. At the 500-m spatial scale, when all detections were combined and compared to random sites, there was no overall difference, but developed land (F = 4.98, P = 0.026) was greater at random sites. Habitat variables differed between detection and random sites for M. grisescens (Wilk?s ? = 0.931, P = 0.015) and P. subflavus (Wilk?s ? = 0.941, P = 0.033). The univariate portion of the analysis and comparison of plots of average area of habitat types indicates that open water (F = 4.34, P = 0.038) and deciduous forest (F = 12.72, P < 0.001) are habitat associations for M. grisescens. Developed land was significantly greater at random sites for M. grisescens (F = 6.14, P = 0.038) and P. subflavus (F = 8.91, P = 0.003), which indicates that these species more often occur outside of these habitats. Wetland (F = 8.78, P = 0.003) was a good predictor of detection of P. subflavus. There was no significant habitat association for E. fuscus, L. borealis, or N. humeralis. At the 1,000-m spatial scale, when all detections were combined and compared to random sites, there was no statistically significant difference and no habitat variable was statistically significant. Habitat variables differed between detection and random sites for M. grisescens (Wilk?s ? = 0.909, P = 0.001) and P. subflavus (Wilk?s ? = 0.943, P = 0.044). Univariate analysis of area for each habitat type indicate that deciduous forest (F = 20.22, P < 0.001) is associated with capture of M. grisescens, and developed land (F = 4.65, P = 0.032) and open pasture (F = 7.14, P = 0.008) were significantly greater at random sites; therefore, M. grisescens occurs less often than expected in these habitats. 43 Developed land (F = 7.86, P = 0.005) also was not a positive habitat association for P. subflavus, but wetland (F = 8.54, P = 0.004) was a habitat association. There was no statistically significant variable for E. fuscus, L. borealis, or N. humeralis. Detection and no-detection sites versus random sites.--There was no statistically significant difference between habitat variables at sites where detectors were placed and random sites for the 250-m buffers. At the 500-m spatial scale, GLM detected a difference between sites where detectors were placed and random sites. Deciduous forest (F = 5.93, P = 0.016) occurred more in sites where detectors were placed. At the 1,000- m spatial scale, there was a difference in amount of developed land between sites where detectors were placed and random sites and developed land (F = 4.98, P = 0.026) occurred more in random sites. Thus, all species of bats occurred less frequently than expected in developed land. DISCUSSION Analyses of data comparing sites where bats were detected with sites with no- detection revealed that detectors that were placed in areas with open water and deciduous forest were significantly more likely to detect bats than detectors that were placed in areas with developed lands. Data from capture versus random sites also indicated that bats were significantly less likely to occur on developed land. As area of developed land increased, bats were less likely to use the landscape for foraging habitat. When data from detection and no-detection sites were compared to random sites, there was no statistically significant difference in composition of habitat types at the 250-m spatial scale. I did not sample randomly at the 500 and 1,000-m scales and chose to place detectors in sites that had more deciduous forest while random sites had more developed land. 44 As expected, I detected no significant habitat association for E. fuscus, which agrees with previous studies that have shown this species is a habitat generalist (Agosta 2002; Kurta and Baker 1990; Chapter 1). An examination of types of habitat where E. fuscus was detected indicated that it occurred at sites with open water, deciduous forest, evergreen forest, mixed forest, open pasture, and wetland. It is clear that E. fuscus is a habitat generalist at Redstone Arsenal. My study demonstrated that L. borealis was associated with open water. This species flies above the tree-canopy and over open pastures (LaVal et al. 1977; Shump and Shump 1982), but had not been reported previously to spend significant time foraging over open water. I determined that M. grisescens foraged over open water and in deciduous forest and avoided developed land and open pasture. M. grisescens forages over open water and in riparian areas using the forest for cover (Best and Hudson 1996; Goebel 1996; Henry 1998; Tuttle 1976), which explains why the species was negatively associated with open areas such as developed lands and open pasture. Although M. grisescens will commute through non-riparian areas on its way to forage in open-water habitat, it is not known to forage in non-riparian areas (Best and Hudson 1996; Tuttle 1979). My study has shown that N. humeralis was associated with open water. N. humeralis forages in woodlands and roosts in buildings (Watkins 1972), but has not been documented to forage over open water. Perimyotis subflavus foraged in open water and wetland habitats and avoided developed land and mixed forest. This species uses streams and forest edges as foraging 45 habitats (Davis and Mumford 1962; Fujita and Kunz 1984; LaVal et al. 1977), but has not been documented to use wetland habitats. Other species of bats that were not detected on Redstone Arsenal, but whose ranges cover the Arsenal are listed in Table 2.3. One of the motives for using ultrasonic detection, as noted in Chapter 1, was that it may be possible to detect species on Redstone Arsenal that were not captured in mist nets. I was particularly interested in detecting Tadarida brasiliensis, a species that flies at high altitudes and is not easily captured in mist nets (Wilkins 1989). I did not detect this species on the Arsenal, and it is difficult to tell whether the species does not occur on Redstone Arsenal or, perhaps, it may occur in the study area but was overlooked by my methods of detection. Further investigation is needed to ascertain the presence of the species here and elsewhere in northern Alabama. Management Implications.--Because I determined that M. grisescens foraged in sites with open water and deciduous forest and because most captures occurred at sites with open water and deciduous forest, I recommend that these areas be preserved for use by bats on Redstone Arsenal. M. grisescens avoids areas with developed land and open pasture, so I encourage facility managers to keep these types of habitats to a minimum when managing for bats. While some studies use results from acoustic detection to determine management plans, other researchers suggest that acoustic detection should be used only for generating a priori hypotheses about use of habitats (Miller et al. 2003). I believe it is necessary to use mist nets to capture bats and have these species in hand for positive identification before finalizing management plans. 46 LITERATURE CITED AEBISCHER, N. J., P. A. ROBERTSON, AND R. E. KENWARD. 1993. Compositional analysis of habitat use from animal radio-tracking data. Ecology 74:1313-1325. AGOSTA, S. J. 2002. Habitat use, diet and roost selection by the big brown bat (Eptesicus fuscus) in North America: a case for conserving an abundant species. Mammal Review 32:179-198. BEST, T. L., AND M. K. HUDSON. 1996. Assessment of routes used by female gray bats (Myotis grisescens) between roost sites and foraging areas in northern Alabama. Journal of the Alabama Academy of Science 67:6-14. BRITZKE, E. R. 2003. Use of ultrasonic detectors for acoustic identification and study of bat ecology in the eastern United States. Ph.D. dissertation, Tennessee Technological University, Cookeville. DAVIS, W. H., AND R. E. MUMFORD. 1962. Ecological notes on the bat Pipistrellus subflavus. American Midland Naturalist 68:394-398. ERICKSON, J. L., AND S. D. WEST. 2003. Associations of bats with local structure and landscape features of forested stands in western Oregon and Washington. Biological Conservation 109:95-102. FENTON, M. B., S. BOUCHARD, M. J. VONHOF, AND J. ZIGOURIS. 2001. Time-expansion and zero-crossing period meter systems present significantly different views of echolocation calls of bats. Journal of Mammalogy 82:721-727. FUJITA, M. S., AND T. H. KUNZ. 1984. Pipistrellus subflavus. Mammalian Species 228:1-6. GLENDELL, M., AND N. VAUGHAN. 2002. Foraging activity of bats in historic landscape 47 parks in relation to habitat composition and park management. Animal Conservation 5:309-316. GOEBEL, A. B. 1996. Temporal variation in movement patterns of adult female Myotis grisescens (Chiroptera: Vespertilionidae). M.S. thesis, Auburn University, Alabama. HENRY, T. H. 1998. Variation in use of habitats by the gray bat (Myotis grisescens) in northern Alabama. M.S. thesis, Auburn University, Alabama. KALCOUNIS, M. C., K. A. HOBSON, R. M. BRIGHAM, AND K. R. HECKER. 1999. Bat activity in the boreal forest: importance of stand type and vertical strata. Journal of Mammalogy 80:673-682. KURTA, A., AND R. H. BAKER. 1990. Eptesicus fuscus. Mammalian Species 356:1-10. LACKI, M. J., S. K. AMELON, AND M. D. BAKER. 2007. Foraging ecology of bats in forests. Pp. 83-127, in Bats in forests: conservation and management (M. J. Lacki, J. P. Hayes, and A. Kurta, eds.). Johns Hopkins University Press, Baltimore, Maryland. LAVAL, R. K., R. L. CLAWSON, M. L. LAVAL, AND W. CAIRE. 1977. Foraging behavior and nocturnal activity patterns of Missouri bats, with emphasis on the endangered species Myotis grisescens and Myotis sodalis. Journal of Mammalogy 58:592- 599. LOEB, S. C., AND J. M. O'KEEFE. 2006. Habitat use by forest bats in South Carolina in relation to local, stand, and landscape characteristics. Journal of Wildlife Management 70:1210-1218. MCCUNE, B., AND J. B. GRACE. 2002. Analysis of ecological communities. MjM 48 Software Design, Gleneden Beach, Oregon. MILLER, D. A., E. B. ARNETT, AND M. J. LACKI. 2003. Habitat management for forest- roosting bats of North America: a critical review of habitat studies. Wildlife Society Bulletin 31:30-44. MURRAY, K. L., E. R. BRITZKE, B. M. HADLEY, AND L. W. ROBBINS. 1999. Surveying bat communities: a comparison between mist nets and the Anabat II bat detector system. Acta Chiropterologica 1:105-112. SAMORAY, S. T. 2002. Bat utilization of barrens restoration sites on Arnold Air Force Base, Tennessee. M.S. thesis, Middle Tennessee State University, Murfreesboro. SHUMP K. A., JR., AND A. U. SHUMP. 1982. Lasiurus borealis. Mammalian Species 183:1-6. TUTTLE, M. D. 1976. Population ecology of the gray bat (Myotis grisescens): factors influencing growth and survival of newly volant young. Ecology 57:587-595. TUTTLE, M. D. 1979. Status, causes of decline, and management of endangered gray bats. Journal of Wildlife Management 43:1-17. VAUGHAN, N., G. JONES, AND S. HARRIS. 1997. Habitat use by bats (Chiroptera) assessed by means of a broad-band acoustic method. Journal of Applied Ecology 34:716-730. WATKINS, L. C. 1972. Nycticeius humeralis. Mammalian Species 23:1-4. WILKINS, K. T. 1989. Tadarida brasiliensis. Mammalian Species 331:1-10. YATES, M. D., AND R. M. MUZIKA. 2006. Effect of forest structure and fragmentation on site occupancy of bat species in Missouri Ozark forests. Journal of Wildlife Management 70:1238-1248. 49 _________ Table 2.1. Significant habitat associations for bats that occur on Redstone Arsenal, Madison Co., Alabama, 2005-2007, as determined by GLM comparing mean area of each habitat type at sites where a species was detected to sites where that species was not detected. A positive sign indicates habitats where bats were detected and a negative sign indicates habitat where bats were not detected. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. ____________________________________________________________________________________________________________ Variable and size of buffer Species Lasiurus borealis Open Water 250-m (+)* Open Water 1,000-m (+)** Myotis grisescens Open Water 250-m (+)** Developed 250-m (-)* Deciduous Forest 250-m (+)** Open Pasture 250-m (-)** Open Water 500-m (+)** Developed 500-m (-)* Deciduous Forest 500-m (+)** Open Pasture 500-m (-)** Deciduous Forest 1,000-m (+)** Open Pasture 500-m (-)** Nycticeius humeralis Open Water 250-m (+)* Perimyotis subflavus Open Water 250-m (+)** Developed 250-m (-)** Mixed Forest 250-m (-)* Open Pasture 250-m (-)* Wetland 250-m (+)** Open Water 500-m (+)** Developed 500-m (-)** Mixed Forest 500-m (-)* Wetland 500-m (+)** Open Water 1,000-m (+)* Developed 1,000-m (-)** Mixed Forest 1,000-m (-)* Wetland 1,000-m (+)* 50 _________ Table 2.2. Significant habitat associations for bats that occur on Redstone Arsenal, Madison Co., Alabama, 2005-2007, as determined by GLM comparing mean area of each habitat type at sites where a species was detected against random sites. A positive sign indicates habitats where bats were detected and a negative sign indicates habitats where bats were not detected. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. ____________________________________________________________________________________________________________ Variable and size of buffer Species Myotis grisescens Open Water 250-m (+)* Deciduous Forest 250-m (+)* Open Pasture 250-m (-)* Open Water 500-m (+)* Developed 500-m (-)* Deciduous Forest 500-m (+)** Open Pasture 500-m (-)* Developed 1,000-m (-)* Deciduous Forest 1,000-m (+)** Open Pasture 500-m (-)* Nycticeius humeralis Open Water 250-m (+)* Perimyotis subflavus Open Water 250-m (+)* Developed 500-m (-)** Wetland 500-m (+)** Developed 1,000-m (-)* Wetland 1,000-m (+)* __________________________________________________________________________________________________ 51 Table 2.3. Species of bats that occur in the geographic range of Redstone Arsenal, Madison Co., Alabama, that were not detected during this study. ___________________________________________________________________________________________________ Corynorhinus rafinesquii Lasiurus cinereus Lasionycteris noctivagans Lasiurus seminolus Myotis austroriparius Myotis leibii Myotis lucifugus Myotis sodalis Myotis lucifugus Tadarida brasiliensis ___________________________________________________________________________________________________ Fig. 2.1. Sites were AnaBat detectors were placed on Redstone Arsenal, Madison, Co., Alabama, summer 2007. 52 Fig. 2.2. Photo of the weather-proof housing in which AnaBat detectors were placed during the study of foraging habitats of bats at Redstone Arsenal, Madison Co., Alabama, 2007. 53 Figure 2.3 Representative echolocation calls for a) Eptesicus fuscus, b) Lasiurus borealis, c) Myotis grisescens, d) Nycticeius humeralis, and e) Perimyotis subflavus as detected by an acoustic-monitoring device at Redstone Arsenal, Madison, Co., Alabama. a) b) c) d) e) 54 Fig. 2.4 Average area (m 2 ) of habitat types at the 250-m spatial scale at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates detection sites and a striped bar indicates no-detection sites for a) Eptesicus fuscus, b) Lasiurus borealis, c) Myotis grisescens, d) Nycticeius humeralis, and e) Perimyotis subflavus. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. 55 Fig. 2.5 Average area (m 2 ) of habitat types at the 500-m spatial scale at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates detection sites and a striped bar indicates no-detection sites for a) Eptesicus fuscus, b) Lasiurus borealis, c) Myotis grisescens, d) Nycticeius humeralis, and e) Perimyotis subflavus. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. 56 Fig. 2.6 Average area (m 2 ) of habitat types at the 1,000-m spatial scale at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates detection sites and a striped bar indicates no-detection sites for a) Eptesicus fuscus, b) Lasiurus borealis, c) Myotis grisescens, d) Nycticeius humeralis, and e) Perimyotis subflavus. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. 57 Fig. 2.7 Average area (m 2 ) of habitat types at the 250-m spatial scale at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates detection sites and a striped bar indicates random sites for a) Eptesicus fuscus, b) Lasiurus borealis, c) Myotis grisescens, d) Nycticeius humeralis, and e) Perimyotis subflavus. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. 58 Fig. 2.8 Average area (m 2 ) of habitat types at the 500-m spatial scale at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates detection sites and a striped bar indicates random sites for a) Eptesicus fuscus, b) Lasiurus borealis, c) Myotis grisescens, d) Nycticeius humeralis, and e) Perimyotis subflavus. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. 59 Fig. 2.9 Average area (m 2 ) of habitat types at the 1,000-m spatial scale at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates detection sites and a striped bar indicates random sites for a) Eptesicus fuscus, b) Lasiurus borealis, c) Myotis grisescens, d) Nycticeius humeralis, and e) Perimyotis subflavus. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. 60 Fig. 2.10 Average area (m 2 ) of habitat types at the a) 250-m, b) 500-m, and c) 1,000-m spatial scales at Redstone Arsenal, Madison Co., Alabama, where a solid black bar indicates sites where detectors were placed and a striped bar indicates random sites. One asterisk indicates P < 0.05 and two asterisks indicate P < 0.01. 61 62 CHAPTER 3: COMPARING SURVEY METHODS TO ELUCIDATE HABITAT ASSOCIATIONS OF BATS ON REDSTONE ARSENAL, ALABAMA ABSTRACT This study compared number of species per night observed using 2 methods of sampling commonly employed to survey for bats; mist nets and ultrasonic detection. A Student t-test was used to compare data obtained using mist nets and ultrasonic-detection devices to assess habitat associations of bats in northern Alabama. Overall, 7 species were captured in mist nets and 6 species were detected ultrasonically. However, more species of bats per night were detected using ultrasonic methods. Unlike other studies, this study revealed that more species were observed using mist nets than ultrasonic- detection devices. This conclusion may be influenced by amount of clutter in habitats that were sampled. INTRODUCTION Two methods of surveying populations of bats include mist nets and ultrasonic detection. Mist nets are a common sampling technique used for capturing bats that allows the observer to examine the animal in-hand to identify species (Kunz and Kurta 1988). Having the bat in hand allows collection of data about population demographics, such as sex, age, and reproductive status, which may be useful in developing management plans (Murray et al. 1999). In the past 2 decades, researchers have used ultrasonic-detection devices for identifying bats (Bell 1980; Britzke 2003; Fenton and 63 Bell 1981; Livengood 2003; Loeb and O'Keefe 2006; O?Farrell 1997; Yates and Muzika 2006). Data collected with mist nets may be biased because they only capture species that fly low and do not capture species that primarily fly over open fields, open water, or above the canopy (Kunz and Brock 1975); ultrasonic-detection devices allow the observer to sample a larger area (O'Farrell and Gannon 1999). Acoustical monitoring also has its biases, namely the under-representation of certain species whose calls are not detected easily or species that fly above the range of the ultrasonic-detection device (Murray et al. 1999). It also is difficult to identify calls of species that were recorded in areas of clutter such as in forested habitats (Jones et al. 2000; Patriquin et al. 2003). The purpose of my study was to compare average number of species detected each night using mist-net surveys and acoustical monitoring. Previous studies have used both sampling techniques at a site to make comparisons between methods (Kuenzi and Morrison 1998; Murray et al. 1999; O'Farrell and Gannon 1999). Because my study focused on sampling as much area of Redstone Arsenal as possible, only one method was used at each site. MATERIALS AND METHODS Data were collected as in Chapters 1 and 2; the study area was described in Chapter 1. I calculated number of species per night determined by mist netting and by ultrasonic detection. A Student t-test was used to determine if there was a significant difference between number of species detected by each method. RESULTS There were 0-5 species captured in mist nets/night and the average was 1.8 species/night (SD = 1.5, n = 59; Table 3.1). There were 0-5 species detected using 64 AnaBat SD1 and the average was 2.6 (SD = 1.6, n = 30). All species captured in mist nets were detected ultrasonically except for the southeastern myotis (Myotis austroriparius). No additional species was identified using ultrasonic detection. There was a statistically significant difference in number of species per night recorded by the 2 methods. Results from the t-test indicate that average number of species per night was greater using ultrasonic detection (P = 0.013, t = 2.55, df = 87). DISCUSSION My results were at variance with findings by others. More individuals were detected using acoustic monitoring than by mist nets; thus, biodiversity was greater when ultrasonic detection was used (Murray et al. 1999; O'Farrell and Gannon 1999). My study documented verification of presence of more species by using mist nets than by using AnaBat detectors, because M. austroriparius was not recorded using ultrasonic detection. Other studies have detected M. austroriparius using AnaBat and reported that it was associated with riparian areas (Britzke 2003; Ford et al. 2006). The reason that I did not detect M. austroriparius using AnaBat detectors is that habitats in which this species occurs are forested and ultrasonic detectors have difficulty detecting calls of species in habitats that have clutter (forested areas; Patriquin et al. 2003). Although I attempted to deploy detectors in areas without clutter, identifications of calls were limited due to clutter. Representation of species on Redstone Arsenal possibly was limited by the effect of clutter (Patriquin et al. 2003). My observations were concordant with those reported by Murray et al. (1999); that is, the endangered gray myotis (M. grisescens) was detected at more sites with AnaBat detectors than with mist nets. This possibly could be because I placed detectors 65 over open water, over which M. grisescens was known to forage, while I could not place nets over large areas of open water. The northern myotis (Myotis septentrionalis) is considered difficult to detect by some researchers (Hickey and Neilson 1995), but has been detected successfully by others (Murray et al. 1999). My study recorded M. septentrionalis for both techniques of monitoring. Mist net surveys provide collection of sex, age, and reproductive condition data that is lost when acoustical detection alone is used (Kuenzi and Morrison 1998). For example, in 1 mist net over a pond, I captured 6 female evening bats (Nycticeius humeralis), of which 5 were either lactating or post-lactating. The site where I placed my mist net was probably near a maternity roost since several female individuals were captured in one net. Use of ultrasonic detection is the least intrusive method and can be used repeatedly at a site while bats will learn to avoid mist nets with repeated encounters at a site (Kunz and Brock 1975). Using both methods together provides a more complete inventory of species than using either method alone because each method has its benefits (Kuenzi and Morrison 1998; O?Farrell and Gannon 1999; Murray et al. 1999). LITERATURE CITED BELL, G. P. 1980. Habitat use and response to patches of prey by desert insectivorous bats. Canadian Journal of Zoology 58:1876-1883. BRITZKE, E. R. 2003. Use of ultrasonic detectors for acoustic identification and study of bat ecology in the eastern United States. Ph.D. dissertation, Tennessee Technological University, Cookeville. 66 FENTON, M. B., AND G. P. BELL. 1981. Recognition of species of insectivorous bats by their echolocation calls. Journal of Mammalogy 62:233-243. FORD, W. M., J. M. MENZEL, M. A. MENZEL, J. W. EDWARDS, AND J. C. KILGO. 2006. Presence and absence of bats across habitat scales in the upper Coastal Plain of South Carolina. Journal of Wildlife Management 70:1200-1209. HICKEY, M. B. C., AND A. L. NEILSON. 1995. Relative activity and occurrence of bats in southwestern Ontario as determined by monitoring with bat detectors. Canadian Field-Naturalist 109:413-417. JONES, G., N. VAUGHAN, AND S. PARSONS. 2000. Acoustic identification of bats from directly sampled and time expanded recordings of vocalizations. Acta Chiropterologica 2:155-170. KUENZI, A. J., AND M. L. MORRISON. 1998. Detection of bats by mist-nets and ultrasonic sensors. Wildlife Society Bulletin 26:307-311. KUNZ, T. H., AND C. E. BROCK. 1975. A comparison of mist nets and ultrasonic detectors for monitoring flight activity of bats. Journal of Mammalogy 56:907- 911. KUNZ, T. H., AND A. KURTA. 1988. Capture methods and holding devices. Pp. 1-29, in Ecological and behavioral methods for the study of bats (T. H. Kunz, ed.). Smithsonian Institution Press, Washington, D.C. LIVENGOOD, K. 2003. The Anabat bat detector's zone of reception and the factors that affect detection volume. M.S. Thesis, University of Missouri-Columbia, Columbia. 67 LOEB, S. C., AND J. M. O'KEEFE. 2006. Habitat use by forest bats in South Carolina in relation to local, stand, and landscape characteristics. Journal of Wildlife Management 70:1210-1218. MURRAY, K. L., E. R. BRITZKE, B. M. HADLEY, AND L. W. ROBBINS. 1999. Surveying bat communities: a comparison between mist nets and the Anabat II bat detector system. Acta Chiropterologica 1:105-112. O?FARRELL, M. J. 1997. Use of echolocation calls for the identification of free-flying bats. Transactions of the Western Section of the Wildlife Society 33:1-8. O'FARRELL, M. J., AND W. L. GANNON. 1999. A comparison of acoustic versus capture techniques for the inventory of bats. Journal of Mammalogy 80:24-30. PATRIQUIN, K. J., L. K. HOGHERG, B. J. CHRUSZCZ, AND R. M. R. BARCLAY. 2003. The influence of habitat structure on the ability to detect ultrasound using bat detectors. Wildlife Society Bulletin 31:475-481. YATES, M. D., AND R. M. MUZIKA. 2006. Effect of forest structure and fragmentation on site occupancy of bat species in Missouri Ozark forests. Journal of Wildlife Management 70:1238-1248. 68 Table 3.1. Values used in a t-test to compare number of species per night that were captured in mist nets and by AnaBat on Redstone Arsenal, Madison Co., Alabama. ___________________________________________________________________________________________________ Number of Species per Night Mist Net AnaBat Range 0 - 5 0 - 7 n 104 94 Mean 1.76 3.13 SD 1.466 2.081 69 APPENDIX 1. ____________________________________________________________________________________________________________ Date, time, general location, GPS coordinates, species, sex, age, and reproductive status of bats captured at Redstone Arsenal, Madison Co., Alabama, 2005-2007. ____________________________________________________________________________________________________________ Time Reproductive Date (CDT) Locality GPS coordinates Species Sex Age status ____________________________________________________________________________________________________________ 20 July 2005 2048 Creek Road 34?38.528?N, Perimyotis Male Adult Testes 86?36.819?W subflavus nonscrotal 2117 Creek Road 34?38.415?N, Myotis Male Young- Immature 86?36.805?W septentrionalis of-Year 2154 Creek Road 34?38.528?N, Myotis Male Young- Immature 86?36.819?W grisescens of-Year 2154 Creek Road 34?38.565?N, Eptesicus Female Adult Postlactating 86?36.809?W fuscus 2201 Creek Road 34?38.528?N, Eptesicus Male Young- Immature 86?36.819?W fuscus of-Year 2326 Creek Road 34?38.528?N, Myotis Male Adult Testes 86?36.819?W septentrionalis nonscrotal 2331 Creek Road 34?38.528?N, Lasiurus Female Young- Immature 86?36.819?W borealis of-Year 2340 Creek Road 34?38.528?N, Perimyotis Male Adult Testes 86?36.819?W subflavus nonscrotal 21 July 2005 0045 Creek Road 34?38.528?N, Lasiurus Female Adult Postlactating 86?36.819?W borealis 0121 Creek Road 34?38.528?N, Perimyotis Female Young- Immature 86?36.819?W subflavus of-Year 0146 Creek Road 34?38.339?N, Lasiurus Male Adult Testes 86?36.869?W borealis nonscrotal 2115 1.25 mile N 34?37.937?N, Eptesicus Female Young- Immature Gate 3 86?35.495?W fuscus of-Year 2138 1.25 mile N 34?37.945?N, Lasiurus Female Adult Postlactating Gate 3 86?35.808?W borealis 2316 1.25 mile N 34?37.945?N, Perimyotis Male Young- Immature Gate 3 86?35.808?W subflavus of-Year 70 APPENDIX 1. Continued. ____________________________________________________________________________________________________________ Time Reproductive Date (CDT) Locality GPS coordinates Species Sex Age status ____________________________________________________________________________________________________________ 22 July 2005 0055 1.25 mile N 34?37.939?N, Perimyotis Male Young- Immature Gate 3 86?35.827?W subflavus of-Year 2053 Recreation 34?35.010?N, Eptesicus Male Young- Immature Area 2 86?36.674?W fuscus of-Year 25 May 2006 2120 S of Shields Road 34?35.045?N Lasiurus Male Adult Testes 86?40.669?W borealis nonscrotal 31 May 2006 2110 S of Patton Road 34?34.532?N Lasiurus Undetermined sex, age, and reproductive status 86?37.882?W borealis 01 June 2006 2220 Indian Creek 34?38.763?N Myotis Female Adult Lactating 86?41.180?W austroriparius 02 June 2006 0120 Indian Creek 34?38.763?N Myotis Male Adult Testes 86?41.180?W austroriparius nonscrotal 2107 Nature Center 34?34.775?N Perimyotis Male Adult Testes 86?37.082?W subflavus nonscrotal 2330 Nature Center 34?34.826?N Myotis Male Adult Testes 86?37.149?W grisescens nonscrotal 10 June 2006 2410 Gate 8 34?41.859?N Myotis Male Adult Testes 86?37.782?W grisescens nonscrotal 14 June 2006 2110 Quarry Pond 34?39.616?N Lasiurus Male Adult Testes 86?38.806?W borealis nonscrotal 15 June 2006 2240 Tupelo Swamp 34?39.070?N Myotis Male Young- Immature on Martin Road 86?37.378?W austroriparius of-Year 16 June 2006 0015 Tupelo Swamp 34?39.044?N Nycticeus Female Adult Lactating on Martin Road 86?37.329?W humeralis 2322 McAlpine Road 34?33.210?N Eptesicus Female Adult Lactating 86?38.803?W fuscus 17 June 2006 2122 Anderson Road 34?41.185?N Eptesicus Female Adult Lactating 71 APPENDIX 1. Continued. ____________________________________________________________________________________________________________ Time Reproductive Date (CDT) Locality GPS coordinates Species Sex Age status ____________________________________________________________________________________________________________ 86?42.448?W fuscus 2149 Anderson Road 34?39.951?N Nycticeus Female Adult Postlactating 86?42.634?W humeralis 2149 Anderson Road 34?39.951?N Nycticeus Female Adult Lactating 86?42.634?W humeralis 2149 Anderson Road 34?39.951?N Nycticeus Female Adult Lactating 86?42.634?W humeralis 2311 Anderson Road 34?41.851?N Eptesicus Female Adult Lactating 86?42.448?W fuscus 2315 Anderson Road 34?40.540?N Eptesicus Female Adult Lactating 86?42.464?W fuscus 2328 Anderson Road 34?39.951?N Nycticeus Female Adult Not pregnant 86?42.634?W humeralis Not lactating 2340 Anderson Road 34?40.121?N Myotis Male Adult Testes 86?42.634?W austroriparius nonscrotal 2354 Anderson Road 34?41.185?N Lasiurus Female Adult Postlactating 86?42.448?W borealis 18 June 2006 0051 Anderson Road 34?41.185?N Lasiurus Female Adult Postlactating 86?42.448?W borealis 0051 Anderson Road 34?41.185?N Lasiurus Undetermined sex, age, and reproductive status 86?42.448?W borealis 0104 Anderson Road 34?39.951?N Nycticeus Female Adult Postlactating 86?42.634?W humeralis 0122 Anderson Road 34?40.540?N Lasiurus Undetermined sex, age, and reproductive status 86?42.464?W borealis 0245 Anderson Road 34?39.951?N Nycticeus Female Adult Postlactating 86?42.634?W humeralis 21 June 2006 2011 East Perimeter 34?38.105?N Lasiurus Female Adult Postlactating 86?36.142?W borealis 2013 East Perimeter 34?38.105?N Lasiurus Female Adult Postlactating 86?36.142?W borealis 2042 East Perimeter 34?38.105?N Nycticeus Male Adult Testes 86?36.142?W humeralis nonscrotal 2042 East Perimeter 34?38.105?N Perimyotis Female Adult Lactating 86?36.142?W subflavus 2042 East Perimeter 34?38.105?N Lasiurus Undetermined sex, age, and reproductive status 86?36.142?W sp. 72 APPENDIX 1. Continued. ____________________________________________________________________________________________________________ Time Reproductive Date (CDT) Locality GPS coordinates Species Sex Age status ____________________________________________________________________________________________________________ 2150 East Perimeter 34?38.743?N Myotis Female Young- Immature 86?36.113?W austroriparius of-Year 22 June 2006 0132 East Perimeter 34?38.105?N Eptesicus Female Adult Lactating 86?36.142?W fuscus 2020 East Perimeter 34?35.158?N Lasiurus Female Adult Postlactating 86?35.234?W borealis 2020 East Perimeter 34?35.158?N Lasiurus Female Adult Postlactating 86?35.234?W borealis 2151 East Perimeter 34?35.197?N Myotis Male Young- Immature 86?35.240?W austroriparius of-Year 2158 East Perimeter 34?35.316?N Perimyotis Male Adult Testes 86?35.186?W subflavus nonscrotal 2210 East Perimeter 34?35.158?N Perimyotis Male Adult Testes 86?35.234?W subflavus nonscrotal 2210 East Perimeter 34?35.158?N Nycticeus Male Adult Testes 86?35.234?W humeralis nonscrotal 2210 East Perimeter 34?35.158?N Myotis Female Adult Lactating 86?35.234?W austroriparius 2324 East Perimeter 34?35.316?N Myotis Male Young- Immature 86?35.186?W austroriparius of-Year 23 June 2006 0120 East Perimeter 34?35.158?N Myotis Female Adult Postlactating 86?35.234?W grisescens 2226 East Perimeter 34?37.883?N Myotis Female Adult Lactating 86?37.828?W austroriparius 12 July 2006 2107 Bradford Sinks 34?36.392?N Eptesicus Female Adult Postlactating 86?42.868?W fuscus 2119 Bradford Sinks 34?37.747?N Eptesicus Male Adult Testes 86?43.004?W fuscus nonscrotal 2127 Bradford Sinks 34?36.016?N Nycticeus Male Adult Testes 86?42.232?W humeralis scrotal 2240 Bradford Sinks 34?37.747?N Eptesicus Male Adult Testes 86?43.004?W fuscus nonscrotal 2307 Bradford Sinks 34?36.392?N Eptesicus Female Adult Postlactating 86?42.868?W fuscus 73 APPENDIX 1. Continued. ____________________________________________________________________________________________________________ Time Reproductive Date (CDT) Locality GPS coordinates Species Sex Age status ____________________________________________________________________________________________________________ 2313 Bradford Sinks 34?36.392?N Eptesicus Female Adult Postlactating 86?42.868?W fuscus 2327 Bradford Sinks 34?36.392?N Lasiurus Male Young- Testes 86?42.868?W borealis of-Year scrotal 2338 Bradford Sinks 34?37.747?N Eptesicus Female Adult Not pregnant 86?43.004?W fuscus Not lactating 13 July 2006 0040 Bradford Sinks 34?37.747?N Eptesicus Female Adult Not pregnant 86?43.004?W fuscus Not lactating 0040 Bradford Sinks 34?37.747?N Eptesicus Female Adult Postlactating 86?43.004?W fuscus 0152 Bradford Sinks 34?36.094?N Eptesicus Female Adult Lactating 86?43.217?W fuscus 0218 Bradford Sinks 34?37.747?N Nycticeus Male Young- Testes 86?43.004?W humeralis of-Year scrotal 0218 Bradford Sinks 34?37.747?N Nycticeus Male Adult Testes 86?43.004?W humeralis scrotal 2300 Test Area 1 34?36.885?N Eptesicus Female Young- Immature 86?39.932?W fuscus of-Year 14 July 2006 0234 Test Area 1 34?36.885?N Lasiurus Female Young- Immature 86?39.932?W borealis of-Year 19 July 2006 2050 S Anderson Road 34?37.263?N Lasiurus Female Young- Immature 86?42.708?W borealis of-Year 2222 S Anderson Road 34?37.263?N Myotis Female Adult Postlactating 86?42.708?W austroriparius 2223 S Anderson Road 34?37.263?N Nycticeus Male Young- Testes 86?42.708?W humeralis of-Year scrotal 20 July 2006 0153 S Anderson Road 34?37.263?N Myotis Female Adult Postlactating 86?42.708?W austroriparius 0204 S Anderson Road 34?37.244?N Nycticeus Male Adult Testes 86?42.937?W humeralis scrotal 0204 S Anderson Road 34?37.244?N Nycticeus Male Adult Testes 86?42.937?W humeralis scrotal 0204 S Anderson Road 34?37.244?N Lasiurus Undetermined sex, age, and reproductive status 86?42.937?W borealis 74 APPENDIX 1. Continued. ____________________________________________________________________________________________________________ Time Reproductive Date (CDT) Locality GPS coordinates Species Sex Age status ____________________________________________________________________________________________________________ 2136 Lady Anne Lake 34?40.304?N Lasiurus Female Adult Postlactaqting 86?42.826?W borealis 2136 Lady Anne Lake 34?40.304?N Lasiurus Male Young- Testes 86?42.826?W borealis of-Year scrotal 2136 Lady Anne Lake 34?40.304?N Nycticeus Male Young Testes 86?42.826?W humeralis of Year scrotal 2136 Lady Anne Lake 34?40.304?N Nycticeus Male Adult Testes 86?42.826?W humeralis scrotal 2136 Lady Anne Lake 34?40.304?N Nycticeus Female Adult Postlactating 86?42.826?W humeralis 2136 Lady Anne Lake 34?40.304?N Nycticeus Female Adult Postlactating 86?42.826?W humeralis 2136 Lady Anne Lake 34?40.304?N Nycticeus Female Young- Immature 86?42.826?W humeralis of-Year 2136 Lady Anne Lake 34?40.304?N Nycticeus Female Adult Postlactating 86?42.826?W humeralis 2136 Lady Anne Lake 34?40.304?N Perimyotis Male Young- Testes 86?42.826?W subflavus of-Year scrotal 2136 Lady Anne Lake 34?40.304?N Eptesicus Male Adult Testes 86?42.826?W fuscus scrotal 2140 Lady Anne Lake 34?40.109?N Nycticeus Female Adult Not pregnant 86?41.886?W humeralis Not lactating 2140 Lady Anne Lake 34?40.109?N Nycticeus Male Adult Testes 86?41.886?W humeralis scrotal 2140 Lady Anne Lake 34?40.109?N Eptesicus Female Adult Postlactating 86?41.886?W fuscus 2140 Lady Anne Lake 34?40.109?N Myotis Female Young- Immature 86?41.886?W grisescens of-Year 2303 Lady Anne Lake 34?40.304?N Lasiurus Female Young- Immature 86?42.826?W borealis of-Year 21 July 2006 0110 Lady Anne Lake 34?40.190?N Eptesicus Female Adult Lactating 86?41.879?W fuscus 0228 Lady Anne Lake 34?40.304?N Lasiurus Female Young- Immature 86?42.826?W borealis of-Year 0228 Lady Anne Lake 34?40.304?N Nycticeus Female Adult Not pregnant 86?42.826?W humeralis Not lactating 2115 Anderson Road 34?37.344?N Lasiurus Male Young- Testes 86?38.306?W borealis of-Year scrotal 75 APPENDIX 1. Continued. ____________________________________________________________________________________________________________ Time Reproductive Date (CDT) Locality GPS coordinates Species Sex Age status ____________________________________________________________________________________________________________ 26 July 2006 2040 Field Training Center 34?40.352?N Eptesicus Female Adult Not pregnant 86?37.023?W fuscus Not lactating 2040 Field Training Center 34?40.352?N Myotis Male Young- Testes 86?37.023?W grisescens of-Year nonscrotal 2040 Field Training Center 34?40.352?N Myotis Undetermined sex, age, and reproductive status 86?37.023?W sp. 2350 Field Training Center 34?40.834?N Eptesicus Male Adult Testes 86?36.780?W fuscus scrotal 27 July 2006 0004 Field Training Center 34?40.352?N Myotis Male Young- Testes 86?37.023?W grisescens of-Year nonscrotal 2110 McKinley 34?35.254?N Lasiurus Female Young- Immature Demolition Range 86?39.389?W borealis of-Year 2140 Lady Anne Lake 34?40.109?N Nycticeus Female Adult Not pregnant 86?41.886?W humeralis Not lactating 2140 Lady Anne Lake 34?40.109?N Nycticeus Male Adult Testes 86?41.886?W humeralis scrotal 2140 Lady Anne Lake 34?40.109?N Eptesicus Female Adult Postlactating 86?41.886?W fuscus 2140 Lady Anne Lake 34?40.109?N Myotis Female Young- Immature 86?41.886?W grisescens of-Year 2303 Lady Anne Lake 34?40.304?N Lasiurus Female Young- Immature 86?42.826?W borealis of-Year 26 July 2006 2040 Field Training Center 34?40.352?N Eptesicus Female Adult Not pregnant 86?37.023?W fuscus Not lactating 2040 Field Training Center 34?40.352?N Myotis Male Young- Testes 86?37.023?W grisescens of-Year nonscrotal 2040 Field Training Center 34?40.352?N Myotis Undetermined sex, age, and reproductive status 86?37.023?W sp. 2350 Field Training Center 34?40.834?N Eptesicus Male Adult Testes 86?36.780?W fuscus scrotal 27 July 2006 0004 Field Training Center 34?40.352?N Myotis Male Young- Testes 86?37.023?W grisescens of-Year nonscrotal 2110 McKinley 34?35.254?N Lasiurus Female Young- Immature Demolition Range 86?39.389?W borealis of-Year 76 APPENDIX 1. Continued. ____________________________________________________________________________________________________________ Time Reproductive Date (CDT) Locality GPS coordinates Species Sex Age status ____________________________________________________________________________________________________________ 2203 McKinley 34?35.612?N Lasiurus Undetermined sex, age, and reproductive status Demolition Range 86?38.647?W borealis 28 July 2006 0156 McKinley 34?35.612?N Lasiurus Female Young- Not pregnant Demolition Range 86?38.647?W borealis of-Year Not lactating 09 May 2007 2042 Recreation Area 2 34?35.028?N, Lasiurus Male Adult Testes 86?36.700?W borealis nonscrotal 2046 Recreation Area 2 34?35.020?N, Eptesicus Female Adult Not pregnant 86?36.665?W fuscus Not lactating 2120 Path to Nature 34?34.932?N, Lasiurus Female Adult Not pregnant 86?37.124?W borealis Not lactating 2129 Recreation Area 2 34?35.020?N, Eptesicus Female Adult Pregnant 86?36.665?W fuscus Not lactating 2129 Recreation Area 2 34?35.020?N, Nycticeus Female Adult Not pregnant 86?36.665?W humeralis Not lactating 0142 Recreation Area 2 34?35.020?N, Lasiurus Male Adult Testes 86?36.665?W borealis nonscrotal 10 May 2007 0042 McAlpine Road 34?33.203?N, Perimyotis Male Adult Testes 86?38.724?W subflavus nonscrotal 11 May 2007 0135 Fitness Trail 34?40.650?N, Eptesicus Male Adult Testes 86?38.327?W fuscus scrotal 0135 Fitness Trail 34?40.650?N, Eptesicus Female Adult Lactating 86?38.327?W fuscus 16 May 2007 0135 Indian Creek and 34?38.674?N, Perimyotis Male Adult Testes Martin Road 86?41.558?W subflavus scrotal 18 May 2007 2100 Weeden Mountain 34?41.187?N, Perimyotis Female Adult Not pregnant 86?38.960?W subflavus Not lactating 23 May 2007 2137 Huntsville Spring 34?37.945?N, Perimyotis Male Adult Testes Branch, Patton Road 86?37.841?W subflavus nonscrotal 24 May 2007 2100 Hansen Road 34?40.623?N, Eptesicus Female Undetermined age and 86?37.150?W fuscus reproductive status 77 APPENDIX 1. Continued. ____________________________________________________________________________________________________________ Time Reproductive Date (CDT) Locality GPS coordinates Species Sex Age status ____________________________________________________________________________________________________________ 0030 Hansen Road 34?40.623?N, Eptesicus Female Adult Lactating 86?37.150?W fuscus 0045 Hansen Road 34?40.623?N, Eptesicus Female Adult Lactating 86?37.150?W fuscus 25 May 2007 0130 N Anderson Road 34?39.735?N, Nycticeus Male Adult Testes 86?42.480?W humeralis scrotal 30 May 2007 2100 Test Area 6 34?41.508?N, Lasiurus Female Adult Pregnant 86?41.973?W borealis 2100 Test Area 6 34?40.679?N, Eptesicus Female Adult Lactating 86?41.974?W fuscus 2100 Test Area 6 34?40.679?N, Nycticeus Male Adult Testes 86?41.974?W humeralis scrotal 0137 Test Area 6 34?41.508?N, Nycticeus Male Adult Testes 86?41.973?W humeralis scrotal 31 May 2007 2225 Bradford Sinks 34?35.871?N, Nycticeus Undetermined sex, age, and reproductive status 86?43.400?W humeralis 1 June 2007 0110 Rock Quarry 34?39.595?N, Lasiurus Undetermined sex, age, and reproductive stataus 86?38.753?W borealis 6 June 2007 2010 Test Area 1 34?36.594?N, Lasiurus Male Adult Testes 86?39.162?W borealis scrotal 7 June 2007 2000 McKinley Range 34?35.589?N, Lasiurus Female Adult Lactating 86?38.962?W borealis 2338 McKinley Range 34?35.589?N, Perimyotis Female Adult Lactating 86?38.962?W subflavus 0145 McKinley Range 34?35.589?N, Nycticeus Male Adult Testes 86?38.962?W humeralis scrotal 8 June 2007 2100 Adams Cave 34?34.277?N, Nycticeus Male Adult Testes 86?37.852?W humeralis scrotal 2211 Adams Cave 34?34.357?N, Eptesicus Male Adult Testes 86?37.864?W fuscus scrotal 2211 Adams Cave 34?34.645?N, Myotis Male Adult Testes 86?37.637?W grisescens scrotal 78 APPENDIX 1. Continued. ____________________________________________________________________________________________________________ Time Reproductive Date (CDT) Locality GPS coordinates Species Sex Age status ____________________________________________________________________________________________________________ 0035 Adams Cave 34?34.645?N, Myotis Male Adult Testes 86?37.637?W grisescens nonscrotal 13 June 2007 2305 Buxton Rd Chemical 34?34.214?N, Lasiurus Female Adult Lactating Contamination Site 86?39.619?W borealis 0136 Buxton Rd Chemical 34?34.214?N, Lasiurus Male Adult Testes Contamination Site 86?39.619?W borealis scrotal 14 June 2007 2050 Water Facility 34?39.936?N, Lasiurus Female Adult Lactating 86?39.445?W borealis 20 June 2007 2117 Creek Road 34?38.597?N, Nycticeus Female Adult Lactating 86?36.833?W humeralis 2117 Creek Road 34?38.597?N, Eptesicus Female Adult Not pregnant 86?36.833?W fuscus Not lactating 2344 Creek Road 34?38.436?N, Myotis Male Young- Testes 86?36.823?W austroriparius of-Year scrotal 0110 Creek Road 34?38.597?N, Myotis Male Adult Testes 86?36.833?W grisescens scrotal 21 June 2007 2100 East Perimeter 34?38.244?N, Myotis Female Adult Lactating 86?36.177?W austroriparius 2100 East Perimeter 34?38.244?N, Myotis Female Young- Not pregnant 86?36.177?W austroriparius of-Year Not lactating 22 June 2007 2106 Timmons 34?35.425?N, Lasiurus Male Adult Testes Cemetery Road 86?36.795?W borealis scrotal 2230 Timmons 34?35.373?N, Eptesicus Male Adult Testes Cemetery Road 86?37.356?W fuscus scrotal 2230 Timmons 34?35.373?N, Eptesicus Female Adult Lactating Cemetery Road 86?37.356?W fusc 2230 Timmons 34?35.373?N, Eptesicus Female Adult Lactating Cemetery Road 86?37.356?W fusc 2359 Timmons 34?35.373?N, Eptesicus Female Adult Lactating Cemetery Road 86?37.356?W fusc 2359 Timmons 34?35.373?N, Eptesicus Female Adult Lactating Cemetery Road 86?37.356?W fusc 0030 Timmons 34?35.373?N, Lasiurus Undetermined sex, age, and reproductive status Cemetery Road 86?37.356?W borealis 79 APPENDIX 1. Continued. ____________________________________________________________________________________________________________ Time Reproductive Date (CDT) Locality GPS coordinates Species Sex Age status ____________________________________________________________________________________________________________ 0128 Timmons 34?35.491?N, Eptesicus Female Adult Not pregnant Cemetery Road 86?37.330?W fuscus Not lactating 27 June 2007 2016 Shield Road 34?35.220N, Lasiurus Female Adult Lactating 86?41.039?W borealis 2025 Shield Road 34?35.220N, Lasiurus Female Adult Lactating 86?41.039?W borealis 12 July 2007 2145 East Perimeter 34?37.933N, Myotis Female Adult Lactating Near Gate 3 86?35.390?W austroriparius 2145 East Perimeter 34?37.934N, Lasiurus Undetermined sex, age, and reproductive status Near Gate 3 86?35.450?W borealis 2240 East Perimeter 34?37.933N, Eptesicus Female Adult Lactating Near Gate 3 86?35.390?W fusc 2340 East Perimeter 34?37.934N, Eptesicus Female Young- Not pregnant Near Gate 3 86?35.450?W fuscus of-Year Not lactating 0040 East Perimeter 34?37.934N, Eptesicus Female Young- Not pregnant Near Gate 3 86?35.450?W fuscus of-Year Not lactating 0040 East Perimeter 34?37.934N, Eptesicus Male Young- Testes Near Gate 3 86?35.450?W fuscus of-Year scrotal 13 July 2007 2100 Frisbee Golf 34?40.914N, Eptesicus Female Adult Not pregnant Course on Vincent 86?38.306?W fuscus Not lactating 2100 Frisbee Golf 34?40.914N, Eptesicus Male Adult Testes Course on Vincent 86?38.306?W fuscus scrotal 2100 Frisbee Golf 34?40.914N, Eptesicus Female Adult Not pregnant Course on Vincent 86?38.306?W fuscus Not lactating 2200 Frisbee Golf 34?41.009N, Lasiurus Male Young- Testes Course on Vincent 86?38.280?W borealis of-Year scrotal 2200 Frisbee Golf 34?40.976N, Myotis Male Adult Testes Course on Vincent 86?38.319?W grisescens scrotal 2300 Frisbee Golf 34?40.863N, Lasiurus Male Young- Testes Course on Vincent 86?38.305?W borealis of-Year scrotal 2300 Frisbee Golf 34?41.009N, Lasiurus Female Young- Not pregnant Course on Vincent 86?38.280?W borealis of-Year Not lactating 2400 Frisbee Golf 34?40.863N, Lasiurus Male Young- Testes Course on Vincent 86?38.305?W borealis of-Year scrotal 0130 Frisbee Golf 34?40.914N, Lasiurus Female Young- Not pregnant Course on Vincent 86?38.306?W borealis of-Year Not lactating 0130 Frisbee Golf 34?40.914N, Eptesicus Male Young- Testes Course on Vincent 86?38.306?W fuscus of-Year scrotal 80 APPENDIX 1. Continued. ____________________________________________________________________________________________________________ Time Reproductive Date (CDT) Locality GPS coordinates Species Sex Age status ____________________________________________________________________________________________________________ 18 July 2007 0015 Madkin Mountain 34?40.212N, Lasiurus Female Young- Not pregnant 86?38.936?W borealis of-Year Not lactating 0057 Madkin Mountain 34?40.212N, Lasiurus Undetermined sex, age, and reproductive status 86?38.936?W borealis 0110 Madkin Mountain 34?40.236N, Eptesicus Male Adult Testes 86?38.830?W fuscus scrotal 19 July 2007 2115 Bradford Mountain 34?35.693N, Perimyotis Female Adult Postlac.tating Rock Shelter 86?42.372?W subflavis 2130 Bradford Mountain 34?36.136N, Lasiurus Male Young- Testes Rock Shelter 86?42.562?W borealis of-Year scrotal 2335 Bradford Mountain 34?36.136N, Eptesicus Female Adult Not pregnant Rock Shelter 86?42.562?W fuscus Not lactating 2335 Bradford Mountain 34?36.136N, Lasiurus Male Young- Testes Rock Shelter 86?42.562?W borealis of-Year scrotal 0040 Bradford Mountain 34?36.136N, Lasiurus Male Young- Testes Rock Shelter 86?42.562?W borealis of-Year scrotal 0100 Bradford Mountain 34?36.136N, Nycticeus Male Adult Testes Rock Shelter 86?42.562?W humeralis scrotal 0145 Bradford Mtn. 34?36.136N, Lasiurus Female Young- Not pregnant Rock Shelter 86?42.562?W borealis of-Year Not lactating 26 July 2007 2215 Pasture Near Gate 1 34?35.243N, Lasiurus Female Young- Not pregnant 86?35.232?W borealis of-Year Not lactating 2335 Pasture Near Gate 1 34?35.243N, Eptesicus Female Adult Not pregnant 86?35.232?W fuscus Not lactating 2358 Pasture Near Gate 1 34?35.243N, Eptesicus Female Adult Not pregnant 86?35.232?W fuscus Not lactating ____________________________________________________________________________________________________________ 81 APPENDIX 2. ____________________________________________________________________________________________________________ Localities where mist nets were placed and no bat was captured at Redstone Arsenal, Madison Co., Alabama, 2005-2007. ___________________________________________________________________________________________________________ Date Locality GPS coordinates 21 July 2005 1.25 mile N Gate 3 34?37.943?N, 86?35.525?W 22 July 2005 Recreation Area 2 34?35.076?N, 86?36.693?W 22 July 2005 Recreation Area 2 34?34.926?N, 87?37.134?W 22 July 2005 Recreation Area 2 34?34.949?N, 87?37.130?W 24 May 2006 34?42.234?N, 86?40.680?W 24 May 2006 Mathew?s Cave 34?42.133?N, 86?41.038?W 24 May 2006 34?42.167?N, 86?40.813?W 24 May 2006 34?41.913?N, 86?41.065?W 25 May 2006 34?35.427?N, 86?40.026?W 25 May 2006 34?35.473?N, 86?40.115?W 25 May 2006 34?35.093?N, 86?40.622?W 26 May 2006 Road to Ponds 48N and 48S 34?35.530?N, 86?36.783?W 26 May 2006 Road to Ponds 48N and 48S 34?35.620?N, 86?36.784?W 26 May 2006 Timmon?s Cemetery Road 34?35.308?N, 86?37.483?W 26 May 2006 Timmon?s Cemetery Road 34?35.389?N, 86?37.353?W 31 May 2006 Adam?s Cave 34?34.652?N, 86?37.735?W 31 May 2006 near Adam?s Cave 34?34.655?N, 86?37.682?W 31 May 2006 South end of Patton Road 34?34.599?N, 86?37.885?W 01 June 2006 Madison Road 34?38.681?N, 86?41.584?W 01 June 2006 Madison Road 34?38.681?N, 86?41.561?W 01 June 2006 Tributary to Indian Creek 34?38.789?N, 86?41.172?W 02 June 2006 Nature Center 34?34.945?N, 86?36.881?W 02 June 2006 Nature Center 34?34.934?N, 86?36.921?W 07 June 2006 Weeden Mountain Road 34?41.035?N, 86?39.016?W 07 June 2006 Weeden Mountain Road 34?41.240?N, 86?38.974?W 07 June 2006 Weeden Mountain Road 34?41.191?N, 86?38.945?W 07 June 2006 Weeden Mountain Road 34?40.886?N, 86?38.851?W 08 June 2006 34?40.592?N, 86?38.225?W 08 June 2006 34?40.643?N, 86?38.349?W 82 APPENDIX 2. Continued. ____________________________________________________________________________________________________________ Date Locality GPS coordinates 08 June 2006 34?40.648?N, 86?38.332?W 08 June 2006 34?40.782?N, 86?38.337?W 09 June 2006 Redstone Links 34?41.954?N, 86?39.909?W 09 June 2006 Redstone Links 34?42.023?N, 86?39.837?W 09 June 2006 Stream near Gate 8 34?41.868?N, 86?37.788?W 14 June 2006 Quarry Pond 34?39.619?N, 86?38.893?W 14 June 2006 Quarry Pond 34?39.534?N, 86?38.801?W 14 June 2006 Quarry Pond 34?39.591?N, 86?38.792?W 15 June 2006 Tupelo Swamp 34?39.768?N, 86?37.236?W 15 June 2006 Tupelo Swamp 34?39.627?N, 86?37.318?W 15 June 2006 Tupelo Swamp 34?38.921?N, 86?37.272?W 16 June 2006 McAlpine Road 34?33.204?N, 86?38.727?W 16 June 2006 Iceberg Lake 34?35.148?N, 86?39.314?W 16 June 2006 Iceberg Lake 34?34.904?N, 86?39.336?W 21 June 2006 East Perimeter 34?37.916?N, 86?36.211?W 21 June 2006 East Perimeter 34?37.945?N, 86?36.176?W 22 June 2006 East Perimeter Creek 34?35.234?N, 86?35.235?W 23 June 2006 East Perimeter 34?37.541?N, 86?37.767?W 23 June 2006 East Perimeter 34?37.560?N, 86?37.804?W 23 June 2006 East Perimeter 34?37.886?N, 86?37.793?W 23 June 2006 East Perimeter 34?37.936?N, 86?37.851?W 28 June 2006 Hansen Road 34?40.676?N, 86?37.177?W 28 June 2006 Hansen Road 34?40.625?N, 86?37.145?W 28 June 2006 Hansen Road 34?40.711?N, 86?37.218?W 28 June 2006 Hansen Road 34?40.329?N, 86?37.148?W 28 June 2006 Hansen Road 34?40.393?N, 86?37.167?W 29 June 2006 Igloo Pond 34?34.047?N, 86?39.396?W 29 June 2006 Igloo Pond 34?34.115?N, 86?39.663?W 29 June 2006 Igloo Pond 34?34.652?N, 86?39.630?W 29 June 2006 Igloo Pond 34?34.847?N, 86?39.580?W 13 July 2006 Test Area 1 34?37.133?N, 86?39.900?W 13 July 2006 Test Area 1 34?36.578?N, 86?39.168?W 83 APPENDIX 2. Continued. ____________________________________________________________________________________________________________ Date Locality GPS coordinates 13 July 2006 Test Area 1 34?36.221?N, 86?39.078?W 14 July 2006 Bradford Mountain 34?35.698?N, 86?42.375W 14 July 2006 Bradford Mountain 34?35.696?N, 86?42.351?W 14 July 2006 Bradford Mountain 34?35.727?N, 86?42.348?W 14 July 2006 Bradford Mountain 34?35.857?N, 86?42.388?W 14 July 2006 Bradford Mountain 34?36.148?N, 86?42.566?W 19 July 2006 South Anderson Road 34?37.378?N, 86?42.832?W 19 July 2006 South Anderson Road 34?37.377?N, 86?42.608?W 20 July 2006 Lady Anne Lake 34?39.475?N, 86?42.008?W 20 July 2006 Lady Anne Lake 34?39.470?N, 86?41.994?W 21 July 2006 Anderson Road 34?37.040?N, 86?39.169?W 21 July 2006 Anderson Road 34?37.579?N, 86?38.395?W 21 July 2006 Anderson Road 34?37.536?N, 86?38.359?W 26 July 2006 Field Training Center 34?40.818?N, 86?37.029?W 26 July 2006 Field Training Center 34?40.363?N, 86?37.045?W 27 July 2006 McKinley Technical and Tactical Demolition Range 34?35.715?N, 86?38.229?W 27 July 2006 McKinley Technical and Tactical Demolition Range 34?35.747?N, 86?38.820?W 27 July 2006 McKinley Technical and Tactical Demolition Range 34?35.596?N, 86?38.938?W 09 May 2007 Path to Nature 34?34.980?N, 86?37.126?W 10 May 2007 McAlpine Road 34?33.229?N, 86?38.876?W 10 May 2007 Igloo Pond 34?34.906?N, 86?39.333?W 10 May 2007 Igloo Pond 34?35.145?N, 86?39.314?W 11 May 2007 Fitness Trail 34?40.593?N, 86?38.231?W 11 May 2007 Fitness Trail 34?40.640?N, 86?38.352?W 11 May 2007 Fitness Trail 34?40.783?N, 86?38.354?W 16 May 2007 Indian Creek and Martin Road 34?38.785?N, 86?41.176?W 16 May 2007 Indian Creek and Martin Road 34?38.758?N, 86?41.182?W 16 May 2007 Indian Creek and Martin Road 34?38.682?N, 86?41.572?W 17 May 2007 Martin Road Tupelo Swamp 34?39.764?N, 86?37.227?W 17 May 2007 Martin Road Tupelo Swamp 34?39.611?N, 86?37.336?W 84 APPENDIX 2. Continued. ____________________________________________________________________________________________________________ Date Locality GPS coordinates 17 May 2007 Martin Road Tupelo Swamp 34?39.058?N, 86?37.333?W 17 May 2007 Martin Road Tupelo Swamp 34?39.073?N, 86?37.377?W 18 May 2007 Weeden Mountain 34?40.888?N, 86?38.851?W 18 May 2007 Weeden Mountain 34?41.022?N, 86?39.010?W 18 May 2007 Weeden Mountain 34?41.241?N, 86?38.969?W 23 May 2007 Huntsville Spring 34?37.541?N, 86?37.770?W Branch, Patton Road 23 May 2007 Huntsville Spring 34?37.579?N, 86?37.802?W Branch, Patton Road 23 May 2007 Huntsville Spring 34?37.886?N, 86?37.792?W Branch, Patton Road 23 May 2007 Huntsville Spring 34?37.887?N, 86?37.824?W Branch, Patton Road 24 May 2007 Hansen Road 34?40.369?N, 86?37.183?W 24 May 2007 Hansen Road 34?40.328?N, 86?37.146?W 24 May 2007 Hansen Road 34?40.707?N, 86?37.212?W 24 May 2007 Hansen Road 34?40.674?N, 86?37.170?W 25 May 2007 North Anderson Road 34?39.254?N, 86?42.755?W 25 May 2007 North Anderson Road 34?39.242?N, 86?42.625?W 25 May 2007 North Anderson Road 34?39.733?N, 86?42.919?W 30 May 2007 Test Area 6 34?41.440?N, 86?41.893?W 30 May 2007 Test Area 6 34?40.123?N, 86?41.757?W 31 May 2007 Bradford Sinks 34?35.804?N, 86?43.358?W 31 May 2007 Bradford Sinks 34?36.206?N, 86?43.210?W 31 May 2007 Bradford Sinks 34?36.222?N, 86?43.135?W 1 June 2007 Rock Quarry 34?39.480?N, 86?38.753?W 1 June 2007 Rock Quarry 34?39.527?N, 86?38.796?W 1 June 2007 Rock Quarry 34?39.617?N, 86?38.888?W 6 June 2007 Test Area 1 34?36.225?N, 86?39.085?W 6 June 2007 Test Area 1 34?36.697?N, 86?39.300?W 6 June 2007 Test Area 1 34?37.130?N, 86?39.901?W 7 June 2007 McKinley Range 34?35.262?N, 86?39.389?W 7 June 2007 McKinley Range 34?35.744?N, 86?38.823?W 85 APPENDIX 2. Continued. ____________________________________________________________________________________________________________ Date Locality GPS coordinates 7 June 2007 McKinley Range 34?35.716?N, 86?38.231?W 8 June 2007 Adams Cave 34?34.631?N, 86?37.761?W 12 June 2007 NW Anderson Road 34?41.049?N, 86?42.880?W 12 June 2007 NW Anderson Road 34?41.264?N, 86?42.542?W 12 June 2007 NW Anderson Road 34?41.397?N, 86?42.446?W 12 June 2007 NW Anderson Road 34?41.566?N, 86?42.440?W 13 June 2007 Buxton Road Chemical 34?34.838?N, 86?39.580?W Contamination Area 13 June 2007 Buxton Road Chemical 34?34.643?N, 86?39.627?W Contamination Area 13 June 2007 Buxton Road Chemical 34?34.041?N, 86?39.692?W Contamination Area 14 June 2007 Water Facility 34?39.937?N, 86?39.445?W 14 June 2007 Water Facility 34?39.900?N, 86?39.230?W 14 June 2007 Water Facility 34?39.778?N, 86?39.100?W 20 June 2007 Creek Road 34?38.635?N, 86?36.796?W 20 June 2007 Creek Road 34?38.366?N, 86?36.819?W 21 June 2007 East Perimeter 34?37.937?N, 86?36.170?W 21 June 2007 East Perimeter 34?38.812?N, 86?36.122?W 21 June 2007 East Perimeter 34?38.872?N, 86?36.124?W 22 June 2007 Timmon?s Cemetery Road 34?35.406?N, 86?36.792?W 27 June 2007 Shield Road 34?35.250?N, 86?40.039?W 27 June 2007 Shield Road 34?35.190?N, 86?40.571?W 27 June 2007 Shield Road 34?35.152?N, 86?40.542?W 11 July 2007 Environmental Area 34?35.308?N, 86?40.088?W Shield Road 11 July 2007 Environmental Area 34?35.287?N, 86?40.086?W Shield Road 11 July 2007 Environmental Area 34?35.303?N, 86?39.974?W Shield Road 11 July 2007 Environmental Area 34?35.340?N, 86?39.986?W Shield Road 12 July 2007 East Perimeter 34?37.145?N, 86?35.281?W Near Gate 3 86 APPENDIX 2. Continued. ____________________________________________________________________________________________________________ Date Locality GPS coordinates 12 July 2007 East Perimeter 34?37.227?N, 86?35.251?W Near Gate 3 18 July 2007 Madkin Mountain 34?40.179?N, 86?38.917?W 18 July 2007 Madkin Mountain 34?40.310?N, 86?38.942?W 19 July 2007 Bradford Mountain Rock Shelter 34?35.701?N, 86?42.359?W 19 July 2007 Bradford Mountain Rock Shelter 34?35.705?N, 86?42.354?W 19 July 2007 Bradford Mountain Rock Shelter 34?35.719?N, 86?42.348?W 20 July 2007 Rustic Lodge 34?34.518?N, 86?37.239?W 20 July 2007 Rustic Lodge 34?34.563?N, 86?37.262?W 20 July 2007 Rustic Lodge 34?34.591?N, 86?37.255?W 20 July 2007 Rustic Lodge 34?34.587?N, 86?37.286?W 25 July 2007 Mathew?s Cave 34?42.123?N, 86?41.036?W 25 July 2007 Mathew?s Cave 34?41.902?N, 86?41.066?W 25 July 2007 Mathew?s Cave 34?41.904?N, 86?41.067?W 25 July 2007 Mathew?s Cave 34?41.790?N, 86?41.097?W 25 July 2007 Mathew?s Cave 34?42.109?N, 86?40.812?W 26 July 2007 Pasture Near Gate 1 34?35.298?N, 86?35.334?W 26 July 2007 Pasture Near Gate 1 34?35.313?N, 86?35.354?W 26 July 2007 Pasture Near Gate 1 34?35.518?N, 86?35.202?W ____________________________________________________________________________________________________________ 87 APPENDIX 3. _________ Localities where AnaBat detectors were placed and bats were detected at Redstone Arsenal, Madison Co., Alabama, 2007. ___________________________________________________________________________________________________________ Date AnaBat ID Locality GPS coordinates Species detected ____________________________________________________________________________________________________________ 09 May 2007 AB1 Path to Nature 34?34.820?N, 86?37.199?W None 09 May 2007 AB2 Path to Nature 34?34.942?N, 86?37.127?W None 09 May 2007 AB3 Recreation Area 2 34?34.706?N, 86?36.941?W Lasiurus borealis Myotis grisecns Permyot ubflavu 10 May 2007 AB4 McAlpine Road 34?33.278?N, 86?38.914?W Eptesicus fuscus Lasiurus borealis Nyctieus humeralis Permyotibfvu 10 May 2007 AB5 Igloo Pond 34?34.875?N, 86?39.377?W Nycticeius humeralis Perimyotis subflavus 10 May 2007 AB6 Igloo Pond 34?35.077?N, 86?39.321?W Nycticeius humeralis 11 May 2007 AB7 Fitness Trail 34?40.780?N, 86?38.359?W None 11 May 2007 AB8 Fitness Trail 34?40.662?N, 86?38.268?W None 11 May 2007 AB9 Fitness Trail 34?40.947?N, 86?38.255?W None 16 May 2007 AB10 Indian Creek and Martin Road 34?38.690?N, 86?41.625?W None 16 May 2007 AB11 Indian Creek and Martin Road 34?38.662?N, 86?41.157?W Perimyotis subflavus 16 May 2007 AB12 Indian Creek and Martin Road 34?38.767?N, 86?41.030?W None 17 May 2007 AB13 Martin Road Tupelo Swamp 34?39.793?N, 86?37.152?W None 17 May 2007 AB14 Martin Road Tupelo Swamp 34?39.509?N, 86?37.486?W None 17 May 2007 AB15 Martin Road Tupelo Swamp 34?38.998?N, 86?37.336?W Lasiurus borealis Nycticeius humeralis Perimyotis ubflavus 18 May 2007 AB16 Golf Course 34?42.013?N, 86?39.788?W None 18 May 2007 AB17 Golf Course 34?41.804?N, 86?39.856?W None 18 May 2007 AB18 Golf Course 34?41.609?N, 86?39.850?W None 23 May 2007 AB19 Huntsville Spring 34?37.562?N, 86?37.779?W Lasiurus borealis Branch, PatonRoad Nycticeius humeralis Perimyotis ubflavus 23 May 2007 AB20 Huntsville Spring 34?38.034?N, 86?37.844?W None Branch, Patton Road 88 APPENDIX 3. Continued. ___________________________________________________________________________________________________________ Date AnaBat ID Locality GPS coordinates Species detected ____________________________________________________________________________________________________________ 23 May 2007 AB21 Huntsville Spring 34?39.030?N, 86?37.749?W Lasiurus borealis Branch, PatonRoad Nycticeius humeralis Perimyotis ubflavus 24 May 2007 AB22 Hansen Road 34?40.416?N, 86?37.166?W Lasiurus borealis Perimyotis subflavus 24 May 2007 AB23 Hansen Road 34?40.658?N, 86?37.174?W Eptesicus fuscus Lasiurus borealis Nyctieus humeralis Permyotibfvu 24 May 2007 AB24 Hansen Road 34?40.377?N, 86?37.297?W None 25 May 2007 AB25 Anderson Road 34?39.235?N, 86?42.829?W Eptesicus fuscus Lasiurus borealis Nycticeius humeralis Perimyotis ubflavus 25 May 2007 AB26 Anderson Road 34?39.241?N, 86?42.575?W None 25 May 2007 AB27 Anderson Road 34?39.417?N, 86?42.633?W None 30 May 2007 AB28 Test Area 6 34?40.165?N, 86?41.879?W Eptesicus fuscus Lasiurus borealis Nycticeius humeralis Perimyotis ubflavus 30 May 2007 AB29 Test Area 6 34?41.464?N, 86?41.978?W None 30 May 2007 AB30 Test Area 6 34?40.383?N, 86?42.053?W None 31 May 2007 AB31 Bradford Sinks 34?35.656?N, 86?43.325?W None 31 May 2007 AB32 Bradford Sinks 34?35.758?N, 86?43.183?W None 31 May 2007 AB33 Bradford Sinks 34?36.090?N, 86?43.223?W Myotis grisescens Perimyotis ubflavus 1 June 2007 AB34 Rock Quarry 34?39.850?N, 86?38.987?W Eptesicus fuscus Lasiurus borealis Myotis grisecns Permyot ubflavu 1 June 2007 AB35 Rock Quarry 34?39.447?N, 86?38.624?W Perimyotis subflavus 1 June 2007 AB36 Rock Quarry 34?39.632?N, 86?38.819?W Eptesicus fuscus Lasiurus borealis Myotis grisescens Nycticeius humeralis Perimyotis subflavus 89 APPENDIX 3. Continued. ___________________________________________________________________________________________________________ Date AnaBat ID Locality GPS coordinates Species detected ____________________________________________________________________________________________________________ 6 June 2007 AB37 Test Area 1 34?38.860?N, 86?40.003?W None 6 June 2007 AB38 Test Area 1 34?38.849?N, 86?40.641?W None 6 June 2007 AB39 Test Area 1 34?38.685?N, 86?42.508?W None 7 June 2007 AB40 McKinley Range 34?35.800?N, 86?38.217?W Eptesicus fuscus Lasiurus borealis Myotis grisecns Nycticeius humeralis Perimyotis ubflavus 7 June 2007 AB41 McKinley Range 34?35.422?N, 86?38.639?W None 7 June 2007 AB42 McKinley Range 34?35.216?N, 86?38.952?W None 8 June 2007 AB43 Adams Cave 34?34.726?N, 86?37.983?W None 8 June 2007 AB44 Adams Cave 34?34.759?N, 86?37.724?W None 8 June 2007 AB45 Adams Cave 34?35.042?N, 86?37.553?W None 12 June 2007 AB46 NW Anderson Road 34?41.035?N, 86?42.662?W Lasiurus borealis 12 June 2007 AB47 NW Anderson Road 34?41.256?N, 86?42.554?W None 12 June 2007 AB48 NW Anderson Road 34?41.460?N, 86?42.433?W None 13 June 2007 AB50 Buxton Road Chemical 34?35.092?N, 86?39.536?W Myotis grisescens Contaminaton Perimyotis subflavus 13 June 2007 AB51 Buxton Road Chemical 34?34.346?N, 86?39.640?W None Contamination 13 June 2007 AB52 Buxton Road Chemical 34?34.056?N, 86?39.883?W None Contamination 14 June 2007 AB53 Water Facility 34?39.940?N, 86?39.521?W None 14 June 2007 AB54 Water Facility 34?39.726?N, 86?39.032?W Lasiurus borealis Myotis grisecns Permyot ubflavu 14 June 2007 AB55 Water Facility 34?39.903?N, 86?39.253?W None 20 June 2007 AB56 Creek Road 34?38.669?N, 86?36.754?W Eptesicus fuscus Lasiurs borealis Myotgiscn is eptrioalis Permyois ubflvu 20 June 2007 AB57 Creek Road 34?38.533?N, 86?36.836?W None 20 June 2007 AB58 Creek Road 34?38.411?N, 86?36.954?W None 21 June 2007 AB59 East Perimeter 34?37.639?N, 86?35.368?W None 90 APPENDIX 3. Continued. ___________________________________________________________________________________________________________ Date AnaBat ID Locality GPS coordinates Species detected ____________________________________________________________________________________________________________ 21 June 2007 AB60 East Perimeter 34?37.925?N, 86?36.187?W None 21 June 2007 AB61 East Perimeter 34?38.727?N, 86?36.125?W Eptesicus fuscus Lasiurs borealis Myotgiscn Perimyotis subflavus 22 June 2007 AB62 Timmon?s Cemetery 34?35.287?N, 86?37.550?W Eptesicus fuscus Lasiurs borealis Myotgiscn 22 June 2007 AB63 Timmon?s Cemetery 34?35.452?N, 86?36.789?W Myotis grisescens Perimyotis ubflavus 22 June 2007 AB64 Timmon?s Cemetery 34?35.525?N, 86?37.371?W None 27 June 2007 AB65 Shield Road 34?35.322?N, 86?41.439?W Lasiurus borealis Myotis grisecns Nycticeius humeralis Perimyotis ubflavus 27 June 2007 AB66 Shield Road 34?35.157?N, 86?40.544?W None 27 June 2007 AB67 Shield Road 34?35.158?N, 86?40.798?W Lasiurus borealis 29 June 2007 AB68 Shield Road/Buxton Road 34?35.598?N, 86?40.208?W None 29 June 2007 AB69 Shield Road/Buxton Road 34?35.759?N, 86?39.885?W None 29 June 2007 AB70 Shield Road/Buxton Road 34?35.459?N, 86?35.551?W Perimyotis subflavus 11 July 2007 AB71 Environmental Area 34?34.934?N, 86?38.899?W None Shield Rad 11 July 2007 AB72 Environmental Area 34?34.578?N, 86?38.789?W Myotis grisescens Shield Rad 11 July 2007 AB73 Environmental Area 34?34.402?N, 86?38.943?W None Shield Rad 12 July 2007 AB74 Environmental Area 34?36.511?N, 86?35.518?W None Shield Rad 12 July 2007 AB75 Environmental Area 34?36.668?N, 86?35.579?W Eptesicus fuscus Shield Rad 12 July 2007 AB76 Environmental Area 34?36.801?N, 86?35.609?W Lasiurus borealis Shield Rad 13 July 2007 AB77 Frisbee Golf Course 34?40.940?N, 86?38.350?W Myotis grisescens Vincet 13 July 2007 AB78 Frisbee Golf Course 34?40.970?N, 86?38.222?W Lasiurus borealis Vincet 13 July 2007 AB79 Frisbee Golf Course 34?41.135?N, 86?38.199?W Myotis grisescens Vincet 18 July 2007 AB80 Madkin Mountain 34?40.135?N, 86?38.849?W Myotis grisescens 91 APPENDIX 3. Continued. ___________________________________________________________________________________________________________ Date AnaBat ID Locality GPS coordinates Species detected ____________________________________________________________________________________________________________ 18 July 2007 AB81 Madkin Mountain 34?40.368?N, 86?39.025?W Myotis grisescens 18 July 2007 AB82 Madkin Mountain 34?40.535?N, 86?39.082?W Myotis grisescens Perimyotis ubflavus 19 July 2007 AB83 Bradford Mountain 34?35.679?N, 86?42.382?W Myotis grisescens Rock Shelter Perimyotis subflavus 19 July 2007 AB84 Bradford Mountain 34?35.699?N, 86?42.331?W Myotis grisescens Rock Shelter Perimyotis subflavus 19 July 2007 AB85 Bradford Mountain 34?36.165?N, 86?42.562?W Lasiurus borealis Rock Shelter Myotis grisescens 20 July 2007 AB86 Rustic Lodge 34?34.494?N, 86?37.272?W None 20 July 2007 AB87 Rustic Lodge 34?34.584?N, 86?37.261?W Lasiurus borealis 20 July 2007 AB88 Rustic Lodge 34?34.643?N, 86?37.277?W None 25 July 2007 AB89 Mathew?s Cave 34?42.104?N, 86?40.998?W None 25 July 2007 AB90 Mathew?s Cave 34?41.954?N, 86?41.081?W Lasiurus borealis 25 July 2007 AB91 Mathew?s Cave 34?41.979?N, 86?40.832?W Eptesicus fuscus Lasiurus borealis 26 July 2007 AB92 Pasture Near Gate 1 34?35.110?N, 86?35.264?W Eptesicus fuscus Lasiurs borealis Myotgiscn is eptrioalis Permyois ubflvu 26 July 2007 AB93 Pasture Near Gate 1 34?35.313?N, 86?35.364?W Perimyotis subflavus 26 July 2007 AB94 Pasture Near Gate 1 34?35.746?N, 86?35.212?W Lasiurus borealis ____________________________________________________________________________________________________________