Night-Roosting Behaviors for the Northern Long- Eared Myotis (Myotis septentrionalis) Under a Bridge Revealed by Time-Lapse Photography Author(s): Keith Geluso, Emma C. Keele, Nicole M. Pauley, Isabella R. Gomez, and Simon P. Tye Source: The American Midland Naturalist, 179(2):287-293. Published By: University of Notre Dame https://doi.org/10.1674/0003-0031-179.2.287 URL: http://www.bioone.org/doi/full/10.1674/0003-0031-179.2.287

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BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Am. Midl. Nat. (2018) 179:287–293 Notes and Discussion Piece Night-Roosting Behaviors for the Northern Long-Eared Myotis (Myotis septentrionalis) Under a Bridge Revealed by Time-Lapse Photography

ABSTRACT.—The northern long-eared myotis (Myotis septentrionalis) occurs across much of eastern North America and is listed as federally threatened in the United States due to pervasive population declines. Limited data are available about roosting behaviors for this imperiled species. We report on night-roosting behaviors for the northern long-eared myotis under a bridge in northwestern Nebraska. Grooming, short visits, and feeding were the most frequently observed behaviors. Grooming, inactivity, and nursing had the longest durations, albeit all averaged ,15 min per event. We also documented movement and urination infrequently. Prey manipulation associated with feeding was a frequent behavior and consisted of individuals facing upward or downward, culling wings, elytra, and legs of large prey items. When facing upward wing and tail membranes formed a cup against the abutment wall that likely limited loss of prey. Individuals used the bridge throughout the night but roosted most frequently at 4 and 8 h after sunset (00:15 and 04:15 h, respectively), with early morning activity dominated by feeding/prey manipulation. Our study showed night roosts were used frequently for many reasons, especially for grooming and consumption of large prey. Our observations represent the first description of night-roosting behaviors for the northern long-eared myotis.

INTRODUCTION The northern long-eared myotis (Myotis septentrionalis) is an insectivorous bat occurring across much of eastern North America, ranging from the Atlantic coast to western Canada and parts of the Great Plains (Hall, 1981; Caceres and Barclay, 2000). The species was common throughout much of its distribution before onset and spread the fungus, Pseudogymnoascus destructans, associated with white-nose syndrome that has killed millions of bats of different species (USFWS, 2013, 2015). Pervasive population declines in eastern parts of its distribution led to the northern long-eared myotis being listed as federally threatened in the United States and as endangered in Canada (Turner et al., 2011; COSEWIC, 2013; Moosman et al., 2013; USFWS, 2015; Ingersoll et al., 2016; Reynolds et al., 2016). The northern long- eared myotis roosts in natural and artificial structures (Whitaker et al., 2006; Ormsbee et al., 2007; Stein and White, 2016); however, limited information is available regarding roosting behaviors, including in anthropogenic structures (Ormsbee et al., 2007). Night roosts serve many important functions for bats including, but not limited to, digestion, energy conservation, feeding, grooming, information transfer, protection from inclement weather or predators, and rest (Kunz, 1982; Ormsbee et al., 2007). Duration of night-roosting bouts vary by species (Ormsbee et al., 2007), and some species can spend up to 75% of their nocturnal activity period at night roosts (O’Shea and Vaughan, 1977). Frequency of night roosting depends on multiple factors, including reproductive status and time of year (Anthony et al., 1981; Barclay, 1982; Ormsbee et al., 2007). Understanding functions of night roosts in anthropogenic structures can benefit conservation efforts toward declining bat species (Ormsbee et al., 2007; Knight and Jones, 2009). Many bat species use anthropogenic structures, such as mines, buildings, and bridges, for night roosts (Ormsbee et al., 2007). The extent to which bats use bridges as night roosts and for what purposes remains unclear for many species (Lewis, 1994; Keeley and Tuttle, 1999; Adam and Hayes, 2000; Ormsbee et al., 2007). Moreover, relatively few studies have quantitatively examined the behaviors at night roosts for many temperate bat species in North America (Ormsbee et al., 2007). Herein we describe night-roosting behaviors of the northern long-eared myotis under a roadway bridge. Use of camera traps with time-lapse photography enabled us to discreetly document frequency and duration of behaviors, details rarely reported from night roosts. Understanding how and when the northern long-eared myotis uses anthropogenic structures will aid in conservation and management strategies, as interactions between this protected species and humans are inevitable. 287 288 THE AMERICAN MIDLAND NATURALIST 179(2)

FIG. 1.—Underside of a small bridge in northern Sheridan County, Nebraska, where camera traps were placed to document night-roosting behaviors of the northern long-eared myotis (Myotis septentrionalis), July 2016. Night roosting was observed in semi- enclosed chambers at the top of both abutments between horizontal and vertical girders. Left and right photographs show chambers on the south and north abutments, respectively. White arrows indicate where individuals roosted on abutments

METHODS We conducted this study at a bridge spanning Larrabee Creek, Sheridan County, in the Pine Ridge region of northwestern Nebraska from 22 to 30 July 2016 (Fig. 1; as the bat species in this study is protected, the specific locality data are available from the corresponding author upon request). The segment of creek by the bridge was generally 2.5 m wide and ,0.5 m deep, bordered by a narrow ribbon of deciduous trees that consisted predominantly of plains cottonwood (Populus deltoides), green ash (Fraxinus pennsylvanica), and willows (Salix). East of the bridge for 0.7 km was a pasture without livestock, therefore tall grasses, forbs, and hydrophytes were abundant on this side of the creek. Horses had heavily grazed vegetation along the creek for about 50 m west of the bridge, and those reaches were more open for bats to drink and fly. The surrounding landscape transitioned to gradual hills and ridges dominated by ponderosa pine (Pinus ponderosa) intermixed with agricultural fields and rangelands for domestic livestock. The bridge was small, with a single span between abutments (Fig. 1). Exposed abutment surfaces under the bridge consisted of horizontally-stacked wooden timbers held in place by seven vertical steel I- beams below one horizontal I-beam. Heights of north and south abutments were about 2 m and 2.5 m above the ground, respectively. The bridge deck (9 3 9 m) was supported by seven steel girders, and its underside was covered by corrugated steel sheet metal. These structural elements resulted in six semi- enclosed areas under the deck, hereafter called chambers. Each chamber was about 1.5 m in width between horizontal girders and 8 m in length between the two bridge abutments. We identified bats as the northern long-eared myotis based on length of ears and size compared to other species captured in the area. In addition we placed forearms bands only on northern long-eared myotis in the region. We observed a number of bats, both solitary and in groups, with forearm bands in photographs to confirm identifications. Individuals were identified as recently volant based on their gray coloration and darker appearance in photographs (see Swier, 2003). These age characteristics originally were learned by capturing bats along an adjacent part of the stream near the bridge. Three camera traps (PC800 HyperFire Professional, Semi-Covert, Reconyx Inc., Holmen, Wisconsin) were deployed in three different chambers under the bridge to document night-roosting behaviors of the northern long-eared myotis with time-lapse photography. No bats were present during daylight hours when installing cameras and downloading images. One camera faced the south abutment and two 2018 NOTES AND DISCUSSION PIECE 289 faced the north abutment (Fig. 1). Cameras were placed in chambers with the greatest quantity of fresh guano and prey remnants. We recovered a sample of prey remnants from below roosting surfaces. We identified prey remnants to order and, if possible, to family and measured remnant lengths. Time-lapse and motion-triggered photographs were collected for nine nights beginning on the evening of 22 July and ending the early morning of 30 July 2016. One camera lost power for about 1.5 nights. Cameras used infrared lighting to illuminate subjects and recorded ambient temperature on photographs. We programed cameras to only record during all dark hours of each night (20:00–06:00 h) from about 15 min before sunset (~20:15 h) to about 15 min after sunrise (~05:37 h). We initially programmed cameras on 22 and 23 July to capture images every 1 min, but we learned that a number of behavioral observations lasted ,1 min. We then adjusted cameras to capture images at 15 s intervals. All cameras also captured 10 additional images in about a 10 s period when motion was detected. This allowed us to examine behaviors in more detail and to more accurately determine duration of behaviors. We saved images in JPEG format with a resolution of 3.1 megapixels (2048 3 1536 pixels). We classified behaviors into seven categories (feeding, grooming, inactivity [rest], movement, nursing, short visits, and urinating) and calculated the frequency of each behavior throughout the study period. Feeding included presence of prey in the mouth of bats, individuals facing upright on abutments, or both behaviors simultaneously. We further classified feeding individuals as either facing upward or downward. Feeding behaviors in the upright position were differentiated from urination by the duration of each activity and presence of a dark streak below the bat after urination. Grooming involved bats continuously licking and cleaning their fur or patagia by twisting their heads, lifting their wings, and maneuvering their uropatagia with frequent scratching. Behaviors were classified as inactive when individuals exhibited negligible movement for .1 min, generally with heads facing downward, ears not perked up, and wings tucked beside the body. We defined movement as bats continually crawling on abutment walls. We distinguished nursing by the presence of a juvenile bat next to an adult with the head under the wing of the mother. We classified short visits as appearances of bats for 1 min without noting another behavior. Visits by bats likely were not independent of each other, as some individuals likely used the bridge multiple times. Although a few individuals had forearm bands, we could not read them and there was no way to determine the total number of bats using the bridge. Each behavioral observation was classified further as either a solitary bat (n ¼ 1) or cluster of individuals (n 2). Instances of mothers with young were counted as a clustering behavior, albeit a single observation. For all other groups of bats, we recorded the behavior of each individual as a unique occurrence. Most night-roosting bouts had a single behavior. For some individuals we observed more than one distinct behavior during a night-roosting bout, and we recorded each behavior separately as different observations. For one night-roosting bout with multiple individuals (three adults and three young) that lasted over an hour with individuals coming and going multiple times, we only recorded the behaviors of adults and each new visit counted as a separate observation. We describe some of the behaviors of young in the results during this one large clustering bout, but they were not counted except when nursing with an adult. We calculated the duration of each behavior with sequential time-lapse and motion-triggered photographs by the time accrued between photographs with bats present. For observations of a single photograph, duration was given a value of 0 sec. We did not determine duration for urination as it always appeared in a single photograph. To plot the total frequency of behaviors throughout the night in relation to sunset, we lumped observations in 1 h bins starting at 20:15, the time of sunset Mountain Daylight Time at Rushville, Nebraska. Each bin included behavioral observations the 30 min before and after each designated hour. We reported descriptive statistics as means 6 standard error (SE).

RESULTS We examined 47,787 images. Nighttime air temperatures under the bridge ranged from 19 to 36 C when bats were present. We did not detect other bat species in chambers under the bridge during this period. The earliest bat observed during the study was ~24 min after sunset (20:39 h), and the latest bat observed was ~28 min before sunrise (05:09 h). We tallied seven behaviors from 180 observational events of the northern long-eared myotis during nine nights. Night-roosting observations predominantly were by solitary individuals (88.3%) with only a few groups of bats (11.7%, 2 to 6 individuals) interacting or clumped together under the bridge. Mothers with nursing young accounted for ,5% of observations. 290 THE AMERICAN MIDLAND NATURALIST 179(2)

FIG. 2.—Frequency of all combined and the three most common (feeding, grooming, and short visits) behaviors throughout the night for the northern long-eared myotis (Myotis septentrionalis) underneath a bridge in northwestern Nebraska. Behaviors were recorded between 20:00–06:00 h (~15 min before sunset to ~20 min after sunrise) from 22 to 30 July 2016. Sunset was at ~20:15 h (0 ¼ 20:15 h, 1 ¼ 21:15 h, 2 ¼ 22:15 h, etc.). Each hour represents the number of observations for each behavior 30 min before and after that time. Sunrise was at ~05:37 h, within the 9th h after sunset

Night-roosting behaviors were dominated by grooming (n ¼ 61), short visits (n ¼ 52), and feeding (n ¼ 41), whereas infrequent behaviors consisted of nursing (n ¼ 9), movement (n ¼ 7), urination (n ¼ 6), and inactivity (n ¼ 4). Behaviors with the longest durations included inactivity (12.8 6 6.3 min), grooming (10.7 6 1.9 min), and nursing (9.7 6 2.2 min), and those lasting ,2 min included movement (1.2 6 0.3 min), feeding (1.2 6 0.2 min), and short visits (0.2 6 0.0 min). Night-roosting behaviors, grouped by each hour past sunset, were least frequent 1 to 2 h past sunset (21:15–22:15 h) and peaked around 4 and 8 h after sunset (around 00:15 h and 04:15 h, respectively; Fig. 2). Feeding was most frequent during the early morning (03:45–05:00 h), whereas grooming and short visits were most frequent about 4 and 8 h after sunset (00:15 and 04:15, respectively; Fig. 2). Nursing events were infrequent and mainly occurred at 3 h (n ¼ 2) and 4 h (n ¼ 4) after sunset. One large clustering event with three adults and three volant young involved individuals huddling close together or atop each other. Adults nursed young and groomed themselves, intermittently leaving and returning to the cluster. Adults, presumably females, also returned to clusters for short visits. Volant young, either solitary or in clusters, mainly groomed themselves or were inactive. After nursing, young would often huddle beneath an adult and remain inactive. Bats maintained a clean roost surface by crawling below the roosting area and facing upward to urinate. Northern long-eared myotis spent 78.0% (n ¼ 32) in the upward position and 22.0% (n ¼ 9) in the downward position while dismantling and consuming prey. When facing upward, wings and tail membranes of bats appeared pressed against the abutment wall, forming a cup-like position. From insect remnants (elytra, wings, and legs) collected below roosts, we classified 53 prey items as Arctiidae 2018 NOTES AND DISCUSSION PIECE 291

(Lepidoptera), Noctuidae (Lepidoptera), Tipulidae (Diptera), or Coleoptera. Length of coleopteran elytra ranged from 9 to 13 mm (11.1 6 1.2 mm, n ¼ 15), Diptera wings ranged from 10 to 27 mm (18.1 6 5.9, n ¼ 19), and Lepidoptera wings ranged from 9 to 34 mm (19.0 6 5.5 mm, n ¼ 19).

DISCUSSION To our knowledge this study is the first description of night-roosting behaviors for the northern long-eared myotis, as rigorous studies on night roosting are inherently difficult with these nocturnal volant organisms (Ormsbee et al., 2007). Night roosts often are described as staging areas dominated by long periods (.1h)of inactivity, some with clustering individuals (Barclay, 1982; Kunz, 1982; Ormsbee et al., 2007; Knight and Jones, 2009). Our data for the northern long-eared myotis showed that generally solitary individuals (88.3%) were active and alert (heads up) at night roosts, with most behavioral events of short duration (10 min). The few clustering behaviors tended to be multiple females with volant young. Duration of night-roosting events varies across species based on their functionality (Kunz, 1982; Ormsbee et al., 2007). Further examination of night roosts for the northern long-eared myotis likely will demonstrate that different types and durations of behaviors vary with season, with longer durations of some behaviors during cooler seasons. Grooming was the most prevalent behavior observed at this night roost. Grooming likely was associated with maintaining flight membranes, flight efficiency and effectiveness, and reducing parasite loads (Kunz, 1982). Other species also groom at night roosts (Anthony et al., 1981; Barclay, 1982). For little brown myotis (Myotis lucifugus), grooming primarily occurs following return from feeding and before nightly departures (Burnett and August, 1981). Grooming appears to have important functionality for bats as it can account for a relatively small percentage (,15%) of their time budget in roosts but accounts for .50% of energy expenditure (Burnett and August, 1981). Frequency of grooming by the northern long-eared myotis also might relate to their feeding behaviors at night roosts. Short visits, another frequent behavior, peaked around 3 to 4 h after sunset, with a less pronounced peak before dawn (Fig. 2). Bats alighted on vertical walls of abutments for bouts lasting 1 min. Bats remained relatively stationary with heads up and facing outward, appearing to observe surroundings. Functionality of this behavior is unclear. Feeding and manipulation of large prey occurred throughout the night with a distinct peak in activity before dawn (Fig. 2). Prey, especially moths, were dismantled underneath the bridge, as it appears that the northern long-eared myotis needs to manipulate large prey before consumption. Small prey likely are consumed while flying, but this species might require night roosts to manipulate large prey that are more energetically profitable. Both our study and that of ter Hofstede et al. (2008) observed bats clinging to a vertical surface with their head facing upward, manipulating prey in their mouth, and using wing and tail membranes as a pouch to catch insect parts as they fed. It is unclear why individuals manipulate prey facing upward or downward, but we suspect it relates to prey size or perceived chance of dropping prey while culling less digestible parts. Other North American species of bats also cull large prey at night roosts (O’Shea and Vaughan, 1977; Lacki and Ladeur, 2001). The upward feeding orientation could be linked to subsequent grooming to clean membranes after prey are dismantled and fall into patagia. Analysis of prey remnants under roosts demonstrated that this species consumed moths, flies, and beetles, some of which were rather large based on remnants below roosts. We documented several infrequent behaviors, including nursing, movement, urination, and inactivity by the northern long-eared myotis under the bridge. Nursing was observed infrequently and had one of the longest durations (~10 min). We would expect this behavior to be more frequent earlier in the summer before young become volant, which likely occurs primarily at day roosts. In our study all young appeared capable of flight and arrived at the night roost without adult assistance. After nursing we observed that young occasionally would huddle beneath the mother and remain inactive. Such a behavior might facilitate thermoregulation or represent rest. Lactation continued after young were volant at the study site, as also has been observed with little brown myotis (Kurta et al., 1989). Another behavior with unknown functionality was constant movement of individuals on abutment walls. Individuals sometimes appeared to smell, taste, or lick the surface, as if seeking some sort of information. We also infrequently observed urination by individuals on abutment walls. The manner in which individuals moved below roosting areas to urinate appears to relate to cleanliness of roost 292 THE AMERICAN MIDLAND NATURALIST 179(2) surfaces, fur, and flight membranes, especially for individuals roosting in clusters. Periods of inactivity (~13 min) rarely were observed, but this behavior had the longest duration. Few publications describe that the northern long-eared myotis uses bridges as night roosts (Ormsbee et al., 2007). Our data demonstrated this species commonly used a bridge for many short-duration behaviors in northwestern Nebraska. Our data might help explain the general lack of night-roosting observations at bridges or other night roosts across its distribution. Remaining at roost sites for short durations limits guano accumulation beneath roosts, making it difficult to detect whether structures are being used as night roosts. Quantity of fecal droppings was rather scant compared to other bridge surveys we have conducted elsewhere (Geluso and Mink, 2009). Conventional methods to survey bridges at night, such as walking under them, likely would flush this species from roosts before detection and identification. We predict northern long-eared myotis might use bridges and other human-made structures as night roosts more frequently than is currently recognized. Use of this bridge might have been elevated because night roosts appeared to be a limited resource on the landscape. Our study area lies along the western edge of the known distribution for the northern long-eared myotis (Caceres and Barclay, 2000). Eastern populations in North America generally inhabit interior of forests (Broders et al., 2006), yet our study site contained open deciduous woodlands that generally were limited to patches and narrow strips along the creek. Limitations of the surrounding natural environment might have contributed to the frequent use of this bridge as a night roost. Such structures might need to be considered in conservation strategies for declining bat species (see Ormsbee et al., 2007; Knight and Jones, 2009). Lastly, our study demonstrates the usefulness of time-lapse photography to examine behaviors of bats at roosts with minimal disturbance to study subjects. Camera traps are self-contained units readily available at moderate costs. We predict use of time-lapse photography for other species will reveal additional behaviors at night roosts, especially short-duration events, even for well-studied species. Short-duration behaviors require short intervals between photographs to accurately approximate durations and occurrences. Use of videography and acoustic recorders would capture greater details at such roosts but likely would be less convenient to deploy, difficult to leave for extended times, and data analysis would be more challenging.

Acknowledgments.—We thank Brett Andersen for assistance in the field, Glen Forney for his generosity for housing while conducting this research in the area, and Kristal Stoner for grant related matters associated with bat research in the Pine Ridge region of Nebraska. Cameras originally were purchased by funds provided by the Nebraska Game and Parks Commission and United States Forest Service on other projects. Travel to the area was funded through the Nebraska Game and Parks Commission, State Wildlife Grants program. This project was completed as part of the Honors Program requirements for students enrolled in the Mammalogy Class at the University of Nebraska at Kearney.

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KEITH GELUSO1, EMMA C. KEELE, NICOLE M. PAULEY, ISABELLA R. GOMEZ, SIMON P. TYE, Department of Biology, University of Nebraska at Kearney, Kearney, Nebraska 68849. Submitted 13 September 2017; accepted 8 January 2018.

1 Corresponding author: Telephone: (308) 865-8982; e-mail: [email protected]