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RESEARCH ARTICLE
Summer day-roost selection by eastern red
bats varies between areas with different land-
use histories
Maria N. Monarchino
1
, Marnie L. Behan
2
, Joseph S. JohnsonID
1
*
1Department of Biological Sciences, Ohio University, Athens, Ohio, United States of America, 2School of
Environment and Natural Resources, The Ohio State University, Athens, Ohio, United States of America
*jjohnson@ohio.edu
Abstract
The eastern red bat (Lasiurus borealis) is widely considered to be in decline, inspiring inter-
est in identifying important habitats for conservation in the eastern United States. Unfortu-
nately, knowledge of important day-roosting habitats is lacking for much of the species’
range. We examined patterns of day-roost selection by male and female eastern red bats at
two study sites in southeastern Ohio, U. S. A, to help fill this information gap. We radio-
tagged 28 male and 25 female bats during the summers of 2016–2019 and located 53 male
and 74 female roosts. Day-roost selection differed between sexes and study areas. In a
mostly even-aged forest with significant historical disturbance, we found males and females
roosting in trees located at higher elevations, with no clear selection based on tree or stand
characteristics. Specifically, males selected trees with larger diameters located at lower,
cooler elevations than females, which selected smaller diameter trees found at higher,
warmer elevations. However, in a forest with less historical disturbance and more structural
diversity, we found sexes differed in how they selected from available habitats. These data
show that heterogeneity in environmental conditions can lead to different patterns in selec-
tion, even between sites located within a small geographic area. They also show that east-
ern red bats sexually segregate on the local landscape in the presence of diverse forest
conditions but may not do so in the absence of such diversity. We recommend managing
forests to maintain structural diversity across an elevational gradient to provide male and
female eastern red bats with suitable day-roosting habitat in southeast Ohio.
Introduction
Successful wildlife conservation relies on accurate descriptions of the habitat features and
resources that species need to survive and reproduce [1]. However, such descriptions may be
difficult to formulate for species with large geographic ranges, as they may occupy a wide
range of environmental conditions and ecological communities [2,3]. Further hindering iden-
tification of important resources, some wide-ranging species also occupy human-altered land-
scapes composed of a mosaic of habitat patches, each with a distinct vegetative composition
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OPEN ACCESS
Citation: Monarchino MN, Behan ML, Johnson JS
(2020) Summer day-roost selection by eastern red
bats varies between areas with different land-use
histories. PLoS ONE 15(8): e0237103. https://doi.
org/10.1371/journal.pone.0237103
Editor: Brock Fenton, University of Western
Ontario, CANADA
Received: March 8, 2020
Accepted: July 19, 2020
Published: August 24, 2020
Copyright: ©2020 Monarchino et al. This is an
open access article distributed under the terms of
the Creative Commons Attribution License, which
permits unrestricted use, distribution, and
reproduction in any medium, provided the original
author and source are credited.
Data Availability Statement: Data are stored and
available on the Dryad Data Repository at:
Johnson, Joseph (2020), Lasiurus borealis day-
roost habitat, Dryad, Dataset, https://doi.org/10.
5061/dryad.5dv41ns45.
Funding: This work was funded by the Ohio
Department of Natural Resources (Project # WAPR
23). The sponsor had no role in study design, data
collection or analysis, decision to publish, or
preparation of the manuscript.
Competing interests: The authors have declared
that no competing interests exist.
and structure that influences habitat use and quality [4,5]. Although this can be a challenge for
conservation, these patches represent opportunities to study how species utilize or fail to utilize
different habitats within the same region, which may help improve management. Additionally,
as the number of factors threating biodiversity continues to increase, improving habitat for
wildlife will remain a priority for conservation [6].
In North America, the eastern red bat (Lasiurus borealis) is a wide-ranging species believed
to be in decline as a result of collisions with commercial wind turbines and habitat loss [7,8].
Several species of bats suffer from fatal collisions at commercial wind-energy facilities, but the
eastern red bat is one of the three most commonly killed species in the Midwest [9,10]. Exact
mortality rates are unclear and vary by year, but an estimated 143,023 eastern red bats were
killed at wind facilities in the United States from 2000 to 2011 [10]. Eastern red bat mortalities
are expected to continue as wind-energy technology becomes more popular and efficient, lead-
ing to concerns over their population viability. While large-scale mortalities at wind-energy
facilities are a newer phenomenon, it has long been assumed that eastern red bats are nega-
tively affected by loss of forest habitat across their range [11,12]. As a species that completely
relies on forests, red bats are responsive to changes in forest communities, and forest manage-
ment conducted with this bat in mind may help slow its decline [13]. Unfortunately, studies of
habitat selection in this species are sparsely distributed across its range, which includes the
entirety of the eastern United States and Canada, as well as parts of Mexico [14].
Forests are central to the ecology of many bat species, with 27 of the 45 species documented
in the United States and Canada known to roost in trees [15]. Day-roosts are important because
they provide bats with shelter from the elements, protection from predators, and a place to raise
young [12,16]. Selection of day-roosts are known to vary widely both within and among spe-
cies. Within species, males and females often select different summer roosts based on their dif-
fering needs and limitations during this time [17,18]. In contrast to males, which often select
cooler roosting environments to facilitate torpor during summer [19,20], females require warm
roosts to help keep thermoregulatory costs as low as possible during gestation and lactation [21,
22]. In species that roost under bark or in tree cavities, males tend to roost more solitarily while
females often select large trees that have space for social groups that significantly increase roost
temperatures and provide energy savings [21–23]. While it is common to see females exhibiting
different roosting strategies from males, such as social roosting, this may vary among species.
Solitary, foliage roosting species, such as the eastern red bat, which hangs from branches and
leaf petioles, lack the benefits of social roosting in both sexes [14]. Because these bats roost in
tree canopies exposed to weather, eastern red bats may select roosts based on temperature to
help compensate for thermally unstable roosting environments [24]. However, there are few
studies that examine the role of temperature and other habitat characteristics in roosting by red
bats, especially considering the large geographic range of this species.
Several characteristics of day-roosts influence roost temperature. Foliage roosting species
have been found selecting roosts that are taller than the height of the surrounding canopy,
which may increase solar exposure and roost temperature [25,26]. Other foliage species such
as the hoary bat (Lasiurus cinereus) choose roost sites that decrease wind exposure, offsetting
convective heat loss [27]. Red bats are also known to select roosts based on characteristics that
have little or no known connection to temperature. For example, red bats have been found
roosting high in mature forest canopies that provide adequate gaps beneath the roost for quick
entry and exit but enough cover to avoid detection by site-oriented predators [12,28]. Addi-
tionally, most studies on eastern red bats have found this species to have an affinity for large
diameter hardwoods [29–31]. However, it is unclear if this roost characteristic is connected to
temperature or some other benefit for foliage roosting bats. Thus, day-roost selection in bats is
influenced by several variables pertaining to habitat at different spatial scales.
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As a result of red bats being influenced by forest structure and composition, forest-manage-
ment regimes are likely to affect habitat selection for this species. For example, in contiguous
forests in Kentucky, red bats roost in mature hardwood stands with high basal area far from
forest edges [31]. In intensively managed forests in the southern United States, these bats roost
within or close to mature hardwood stands with a closed canopy [26] or in pine stands that
retained mature hardwoods [32]. While these studies provide insight as to how bats utilize
managed forests, many forests are not intensively managed, but are left to regenerate following
disturbance. This results in different forest structures and possibly different patterns of habitat
use. This is true of many forests in the Western Allegheny Plateau Ecoregion [33], where not
only have forests been heavily disturbed and remain unmanaged, but also where there is cur-
rently no local information on roosting behaviors of the eastern red bat. Determining habitat
characteristics important to eastern red bats in this region may help local management agen-
cies to promote or protect habitat for this declining bat species.
The purpose of this study was to compare day-roost selection by male eastern red bats to
that of females in southeastern Ohio forests with markedly different land use histories and for-
est structures. We used an information theoretic approach to rank a suite of a priori models
seeking to explain day-roost selection within each site. This approach allowed for a compari-
son of top models of selection in each study location to determine if patterns in selection differ.
We hypothesized that (i) eastern red bats would select day-roosts non-randomly, with different
habitat characteristics used by males and females, and (ii) that selection patterns would differ
between sites with different forest conditions.
Materials and methods
Approval for research was given by the Institutional Animal Care and Use Committee of Ohio
University (17-H-008) and the necessary permits (Endangered Species Recovery Permit #
TE5674B) were acquired before conducting field work. All study methods followed the guide-
lines of the American Society of Mammalogists [34].
Study areas
Our study was conducted at two locations in Hocking County, Ohio, each with different land
use histories and forest communities. Our first site, Crane Hollow Nature Preserve (N
39.480089, W -82.584173), is approximately 768 hectares of state nature preserve. Crane Hol-
low Preserve is a biologically diverse area with 10,000 documented floral and faunal species
inhabiting the ridgetops and Blackhand Sandstone gorges that run the length of the preserve.
Elevation at this location ranges from 335 m on ridges to 228 m at the bottom of the gorges.
The forest community along ridgetops is dominated by oak (Quercus spp.), maple (Acer spp.),
hickory (Carya spp.), and elm species (Ulmus spp.), as well as American beech (Fagus grandifo-
lia) and tulip poplar (Liriodendron tulipifera). Eastern hemlock (Tsuga canadensis), American
sycamore (Platanus occidentalis), American beech, birch (Betula sp.), tulip poplar, and maple
species dominate the steep slopes and bottoms of the gorges. Although extensive farming and
timber harvesting has occurred across the preserve before the property became a state preserve
in 1977, large portions of Crane Hollow Preserve have maintained stands that are estimated to
be over 100 years old.
Our second site, located on 1348 hectares owned by the Ohio Division of Wildlife approxi-
mately 33 km northeast of Crane Hollow Preserve, is a section of O’Dowd Wildlife Area locally
referred to as Sunday Creek Coal (N 39.531083, W -82.198662). From the late 1800’s to the
1950’s, Sunday Creek Coal was the location of large coal mining operations that, along other
industry activities, left the property deforested. Much of the land has naturally reforested since
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the closing of the final mine in the 1950’s, although disturbances such as timber harvests, oil
and natural gas exploitation, and the creation of extensive ATV trails are common. The topog-
raphy at Sunday Creek Coal is characterized by hillsides built up by surface mining. Slopes are
not as precipitous as Crane Hollow Preserve and lacked exposed cliffs, but elevations were sim-
ilar, ranging from 222 to 323 m. The forest community consists of a variety of tree species
commonly found in Ohio, such as oaks maple, tulip poplar, bigtooth aspen (Populus grandi-
dentata), American beech, American sycamore, and elms species. Due to the recent mining
operations, stands at Sunday Creek Coal tended to be more densely stocked with smaller,
younger stems compared to those at Crane Hollow Preserve.
Capture and radio-telemetry
We captured bats from late May through early August 2016–2019 using mist nets (Avinet Inc,
Portland, Maine) placed in foraging corridors such as service roads, streams, or over open
water sources. We opened nets shortly before sunset and left them open for 5 hours. All cap-
tured bats were identified to species, age (adult or young-of-the-year), sex, and reproductive
condition. Right forearm length and weight were also measured. Eastern red bats weighing
>6.6 g were fitted with 0.33 g transmitters (model LB-2 and LB-2TX, Holohil Systems Ltd.,
Ontario, Canada) using surgical adhesive (Osto-bond, Quebec, Canada). We tracked radio-
tagged bats to their day-roosts using handheld receivers (Biotracker Version, Lotek Wireless,
Ontario, Canada) each day for the life of the transmitter or until the transmitter had fallen off.
We recorded the UTM of each day-roost using a handheld GPS unit with a 3 m accuracy.
Day-roosting habitat
To compare habitats used to those available, we collected a suite of measurements at the tree-,
stand-, and landscape-scale at each day-roost and at 100 random trees in each study area.
Tree-scale habitat measurements included tree species, diameter at breast height (DBH), tree
height, percent canopy cover, and slope aspect. Stand-scale measurements included canopy
height, distance to nearest live tree, percent slope, and basal area. Landscape-scale measure-
ments included distance to roads, distance to water, distance to forest edge, and elevation.
Basal area was measured using a 10-factor wedge prism, height was estimated with a laser ran-
gefinder/hypsometer, percent canopy cover was determined using a spherical densitometer,
and DBH was measured using a DBH tape. Landscape-scale measurements were calculated
using ArcMap [35].
We generated random locations in each study area to measure habitat available for day-
roosting. Before generating these locations, we determined the area covered by dominant habi-
tat types within each study site and dispersed 100 points proportionally within each of those
habitats. Dominant habitat types were provided by previous studies [36] and from the Ohio
Department of Natural Resources. At Crane Hollow Preserve, these habitats were oak/mesic
forests, hemlock forests, and forested floodplains. At Sunday Creek Coal, the dominant habitat
types were xeric forests, mesic forests, floodplain forests, and a small amount of unclassified
habitat. Coordinates for random points were created using a random point generator in Arc-
Map. We navigated to these locations using a handheld GPS and collected the same suite of hab-
itat variables described above for a tree >10 cm in diameter located closest to the random point.
Ambient temperatures
To investigate if day-roost selection is influenced by ambient air temperatures (T
a
), we mea-
sured T
a
across the elevational gradient present in each study area. We deployed temperature
and relative humidity dataloggers (Onset MX2301, Onset Computer Corp., Bourne,
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Massachusetts) inside solar radiation shields at three slope positions. These corresponded to
ridgetops (Crane Hollow Preserve: 334 m; Sunday Creek Coal: 323 m), mid-slopes (288 and
270 m, respectively), and bottoms (228 and 222 m, respectively). Each logger was programmed
to record T
a
every ten minutes. Dataloggers were deployed in wooded areas and affixed to
small diameter trees approximately 1 m from the ground. Because we were interested in how
temperatures varied across slope positions during the day-roosting period, we summarized T
a
recorded at each logger during the hours between sunrise and sunset into daily average tem-
peratures. In addition, we further limited our summary to data collected during the summer
(June–August) of 2018.
Data analysis
Male and female eastern red bat capture rates were calculated for each site by dividing the total
number of each sex captured by capture effort (nights of netting) to provide a measure of capture
success. We did not statistically compare capture rates as mist-netting is not a reliable technique
for measuring abundance [37], but we do present capture rates as a qualitative measure of capture
success. To determine if forest conditions differed between our two study areas, we conducted a
two-sample t-test on forest measures such as basal area, DBH, tree height, and number of stems
within 20 m from our random trees at both sites. In addition, we conducted a two-sided Fishers
exact test of independence between our female, male, and available trees at both sites to determine
if bats were using certain species more than expected. Similarly, we used a Fisher’s exact test to
determine whether available tree species differed between the two study locations.
To test our two hypotheses, we compared the habitat characteristics of male roosts, female
roosts, and random trees at each site using multinomial logistic regression and a model selec-
tion approach. We used roost trees as the experimental unit and assumed all trees to be inde-
pendent. To assess if using trees, for which we gathered an unequal sample size among bats, as
the experimental unit biased our analyses, we generated average roost characteristics for each
bat and compared the distribution of these average values to that of all trees (data not shown).
These distributions were similar, and we used individual trees in our analysis. Prior to analysis,
we tested for multicollinearity in RStudio [38] using a correlation matrix and did not include
variables in a model if they were >70% correlated. We ran 20 multinomial logistic regression
models for each study area, with each model representing a distinct, biologically informed a pri-
ori model developed from a review of existing literature (S1 Table). Our dependent variable was
sex, which was organized into three categories (male, female, and random trees) to compare
roosts of each sex to available habitat. To assess our hypothesis that day-roost selection would
differ between the two areas, we ranked models for each study area separately using Akaike’s
Information Criterion corrected for small sample sizes (AICc). Models with ΔAICc 2 were
considered competitive as a top model [39]. Using this model selection approach, we compared
our top models from each study location to assess our hypothesis that habitat selection would
differ between the two study areas. We assessed the fit of our top models using Hosmer-Leme-
show goodness of fit tests and referenced our odds ratios to see which variables were driving
our top models. Confidence ratios for our odds ratios were also assessed and any variable with
confidence intervals crossing 1 were not considered to be a strong predictor.
Results
Capture and radio-telemetry
We captured bats on 69 nights, including 39 nights at Crane Hollow Preserve and 30 nights at
Sunday Creek Coal. Adult males were more commonly captured than adult females at both
sites. We captured 1.1 adult male and 0.6 adult female red bats/night at Crane Hollow Preserve
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(n= 43 male, 22 female), and 0.9 adult male and 0.2 adult female bats/night captured at Sunday
Creek Coal (n= 28 male, 7 female). We radio-tagged 17 adult females and 14 adult males at
Crane Hollow Preserve, tracking these bats to 89 roosts (male: n= 29, female: n= 60). At Sun-
day Creek Coal, we radio-tagged 7 adult females and 16 adult males, tracking them to 51 roosts
(male: n= 34, female: n= 17). Overall, bats were tracked for an average of 6.5 days (range =
1–19) to an average of 3.3 day-roosts (range = 1–11) per bat. Bats were captured in the highest
density during the months of June and July at both of our study locations. Males were most
captured during June (n = 38) and July (n = 40), while females had the highest numbers of cap-
tures in June (n = 14). Pregnant bats were detected throughout May (n = 5) and June (n = 7),
while lactating bats appeared (n = 4) in June and continued into July (n = 5). Post-lactating
females (n = 4) were detected from mid- to late-July.
Day-roosting and available habitat
Eastern red bats switched roosts frequently at both study areas. On average, bats switched
roost trees every 1.3 days (range = 1–3) at Sunday Creek Coal and every 1.1 days (range = 1–14)
at Crane Hollow Preserve. At Sunday Creek Coal, females switched every 1.0 days and males
switched every 1.1 days (range = 1–3). Female bats at Crane Hollow Preserve switched roosts
every 1.4 days (range = 1–14) and males switched every 1.1 days (range = 1–3). Habitat mea-
surements from our random trees indicated a significantly different forest structure and com-
position between the study areas. Mean basal area at Sunday Creek Coal was 16.3 ±0.86 m
2
/ha
compared to 23.0 ±1.1 m
2
/ha at Crane Hollow Preserve (t(189) = 4.87, p >0.001). Tree diam-
eter averaged 30.4 ±1.2 cm at Sunday Creek Coal compared to 37.8 ±5.2 cm at Crane Hollow
Preserve (t(184) = 3.84, p <0.001), and average tree height was 18.7 ±0.47 m at Sunday Creek
Coal compared to 20.5 ±0.67 cm (t(178) = 2.25, p = 0.03). Finally, number of live stems within
20 m at Sunday Creek Coal was 69.8 ±4.6 compared to 51.9 ±3.0 stems at Crane Hollow Pre-
serve (t(195) = -4.74, p <0.001).
The species composition of the 100 random trees sampled at each site reflect different forest
communities (p <0.001). At Crane Hollow Preserve, there was a significant difference
between roost tree and available tree species on the landscape (p <0.001). The most frequently
used species were sugar maple (n = 29), oak species (n = 20), and tulip poplar (n = 14). Inspec-
tion of our residual values shows that for females, sugar maple was used significantly more
than expected (Table 1). At Sunday Creek Coal, the residuals also suggest a trend towards
greater use of sugar maple and sweet birch than expected for females, but the omnibus tests
did not find a significant difference between roost and available trees on the landscape
(p = 0.4163). The most frequently used tree species were tulip poplar (n = 13), oak species
(n = 10), sugar maple (n = 8), and American beech (n = 8).
The best supported model from Crane Hollow Preserve was the Tree Diameter and Eleva-
tion Model, which received 69% of the overall model support (Table 2). No other model had
ΔAICc 2 [40]. Odds ratios of variables in this model show that the variable with the largest
impact on probability of a tree being used by females was elevation (odds ratio = 1.151,
CI = 1.151–1.153), but less so for males (odds ratio = 0.986, CI = 0.98–0.99). These ratios show
that for every meter increase in elevation, the likelihood that a bat would occupy a roost
increased by 15% for females but decreased by 1% for males (Fig 1). Females roosted within a
narrow range of elevations (�
x= 330 ±0.07 m, range = 319–334 m) while males (�
x= 319 ±0.41
m, range = 251–331) and random trees (�
x= 321 ±1.46 m, range = 282–335) were found across
a wider range (Fig 2). For males, the variable with the largest effect on probability of a tree
being used was DBH (odds ratio = 1.05, CI = 1.027–1.083). This indicates that for every centi-
meter increase in diameter of a tree, the odds that a male will use that tree for a roosting
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location increases by 5% (Fig 1). However, diameter was not as informative as elevation in
driving female selection (odds ratio = 1.02, CI = 1.002–1.05, Fig 1). Mean diameter of male
(�
x= 52 ±0.52 cm), female (�
x= 44 ±0.30 cm), and random trees (�
x= 38 ±1.55 cm) reflect
these odds ratios (Fig 3).
Table 1. Standardized Pearson’s residuals comparing roost tree species used by male, female, and available trees
on the landscape in two study areas in Ohio from 2016–2019.
Species Female Male Available
Crane Hollow Preserve
Acer rubrum -1.592 -0.282 1.385
Acer saccharum 2.417 -0.289 -1.716
Betula alleghaniensis 0.458 1.251 -1.029
Betula lenta -0.656 1.125 -0.098
Carya spp. 0.794 -0.959 -0.098
Fagus grandifolia -0.656 -0.959 1.025
Fraxinus spp. 0.049 0.795 -0.466
Juglans nigra 1.713 -0.554 -1.029
Liriodendron tulipifera -0.088 1.007 -0.474
Oxydendrum arboretum -0.563 -0.392 0.647
Pinus virginiana -1.127 -0.783 1.295
Platanus occidentalis 0.049 -0.678 0.328
Prunus serotine -0.797 -0.554 0.916
Quercus spp. 0.578 1.394 -1.198
Robinia pseudoacacia -0.563 -0.392 0.647
Sassafras albidum 1.211 -0.392 -0.727
Tsuga canadensis -2.52 -1.752 2.895
Ulmus spp. 0.522 0.893 -0.885
Sunday Creek Coal
Acer rubrum 0.007 -1.41 0.807
Acer saccharinum -0.332 -0.47 0.405
Acer saccharum 2.004 -0.583 -0.479
Ailanthus altissima -0.332 -0.47 0.405
Betula lenta 2.678 -0.47 -0.818
Carya spp. 0.415 -0.282 -0.006
Fagus grandifolia -0.7 1.518 -0.588
Fraxinus spp. -0.47 -0.664 0.573
Liquidambar styraciflua -0.332 -0.47 0.405
Liriodendron tulipifera 1.086 0.329 -0.63
Nyssa sylvatica 0.84 0.124 -0.413
Ostrya virginiana 1.162 0.415 -0.711
Pinus virginiana -0.332 -0.470 0.405
Platanus occidentalis -0.94 -1.329 1.145
Populus grandidentata -0.814 0.587 -0.006
Prunus serotine -0.664 -0.94 0.81
Quercus spp. -1.27 0.923 -0.014
Robinia pseudoacacia -0.575 0.415 -0.005
Ulmus spp. 0.743 0.853 -0.188
Values -2 and 2 are considered influential and are in bold. Positive values indicate greater influence than
expected and negative values indicate less influence than expected.
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In contrast to Crane Hollow Preserve, the best supported model for Sunday Creek Coal was
the Elevation and Distance to Water Model, which received 72% of the overall model support
(Table 3). No other model had ΔAICc 2. Odds ratios of variables in this model show that ele-
vation was the variable with the largest impact on the probability of a tree being used by both
males (odds ratio = 1.05, CI = 1.02–1.077) and females (odds ratio = 1.034, CI = 1.025–1.077).
Thus, each meter increase in elevation had a lesser effect on the probability of a tree being used
by females compared to Crane Hollow Preserve (3% versus 15% increase) but a greater effect for
males (1% decrease versus 5% increase, Fig 1). As a result, males (�
x= 297 ±0.72 m, range =
285–304) and females (�
x= 294 ±0.59 m, range = 251–331) occupied similar ranges of elevations
at Sunday Creek Coal. Random trees were found across a wider range of elevations (�
x=
283 ±1.71 m, range = 242–319) (Fig 2). Distance to water was an explanatory variable in the
model for both males (odds ratio = 1.005, CI = 1.0002–1.0009) and females (odds ratio = 1.009,
CI = 1.003–1.015, Fig 1), with both sexes roosting farther from water sources (females: �
x=
204 ±3.22 m, males: �
x= 164 ±3.66 m) compared to random trees (�
x= 120 ±9.40 m).
Ambient temperatures
Ambient temperatures varied across the elevational gradient at both sites. Ridgetops had slightly
warmer average daytime temperatures from June–August at Crane Hollow Preserve
Table 2. Top 4 models for day-roost selection by male and female eastern red bats, along with the number of
parameters, AICc score, ΔAICc score, and model weight for each, for Crane Hollow Preserve.
Model K AICc Δi wi
Diameter + Elevation 4 334.15 0 0.686
Elevation + Stand 6 336.97 2.81 0.245
Tree Size Model 1 + Elevation 5 337.71 3.55 0.116
Elevation + Distance to Water 4 340.62 6.46 0.027
AIC scores <2 indicates top model.
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Fig 1. Forest plot representing the odds ratios for each variable included in top models for male and female
eastern red bats at Crane Hollow Preserve (top panel) and Sunday Creek Coal (bottom panel) study areas in
southeast Ohio.
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Fig 2. Box and whisker plot showing male (light grey) and female (white) eastern red bats roosting at different
elevations than randomly sampled trees (dark grey) at Crane Hollow Preserve and Sunday Creek Coal study areas
in southeast Ohio, 2016–2019. Females roosted at higher elevations than males at the preserve, but sexes roosted at
similar elevations at the reforested mine. Medians are depicted by the solid line, means are represented by the X’s, and
outliers are depicted as black dots. Quartiles 2 and 3 are represented by the box and quartiles1 and 4 are represented
by whiskers.
https://doi.org/10.1371/journal.pone.0237103.g002
Fig 3. Box and whisker plot showing roost tree diameter of male (light grey) and female (white) eastern red bats at
Crane Hollow Preserve and Sunday Creek Coal study areas in southeast Ohio, 2016–2019. Males roosted in larger
diameter trees while the diameter of female roosts was similar to that of randomly sampled trees. Medians are depicted
by the solid line, means are represented by the X’s, and outliers are depicted as black dots. Quartiles 2 and 3 are
represented by the box and quartiles 1 and 4 are represented by whiskers.
https://doi.org/10.1371/journal.pone.0237103.g003
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(�
x= 24.0 ±0.25˚ C) and Sunday Creek Coal (�
x= 22.6 ±0.22˚ C) compared to lower elevations (�
x=
21.0 ±0.16˚ C and �
x= 20.8 C ±0.15˚ C, respectively). Similar elevations between the two sites varied
slightly in temperature with Crane Hollow Preserve being slightly warmer on average (Fig 4).
Discussion
We found that daytime habitat selection by eastern red bats in southeast Ohio varied between
sexes and between study areas with different forest conditions. In a forest characterized by
Table 3. Top 4 models for day-roost selection by male and female eastern red bats, along with the number of
parameters, AICc score, ΔAICc score, and model weight for each, for Sunday Creek Coal.
Model K AICc Δi wi
Elevation + Distance to Water 4 242.04 0 0.720
Tree Size Model 1 + Elevation 5 245.75 3.71 0.113
Slope Aspect + Elevation 5 246.67 4.64 0.070
DBH + Elevation 4 248.07 6.03 0.035
AIC scores <2 indicates the top model.
https://doi.org/10.1371/journal.pone.0237103.t003
Fig 4. Average daily ambient temperatures were consistently warmer at the highest local elevations present at our
two study areas, Crane Hollow Preserve (panel A) and Sunday Creek Coal (panel B) southeast Ohio from 20 June–
29 August 2018. The dark, dashed, and gray lines represent temperatures collected from the bottom, middle, and
upper elevations sampled.
https://doi.org/10.1371/journal.pone.0237103.g004
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larger, less densely stocked trees, males and females selected day-roosts differently based on
tree diameter and elevation. Female use was more affected by elevation while male use was
more affected by tree diameter. Female roosts were located at higher elevations and in smaller
diameter trees than males. However, in a forest with a similar elevational gradient character-
ized by a higher density of smaller trees, tree diameter was not important in roost selection,
and male and female roosts overlapped more in elevation. These data show that patterns of
habitat use by bats can vary between areas located in the same part of a species’ range, likely in
response to forest structure and microclimate. This highlights the need to study habitat use
across a range of environmental conditions to untangle local effects on habitat use.
The possibility that heterogeneity in environmental conditions can lead to differing effect
sizes, and even directions, for variables influencing habitat selection in bats was recently
explored by Fabianek and colleagues [41]. Focusing solely on cavity and bark roosting species,
the authors found both consistent and inconsistent trends for the influence of variables mea-
sured in the present study, and concluded that summer temperatures moderate some, but not
all, of this variability. The importance of temperature for small, insectivorous bats is well-
known, as these small mammals lose heat quickly, and reproductive females benefit from
selecting day-roosts that minimize this heat loss [23,42]. Warmer maternity roost tempera-
tures are also essential for juvenile development, as offspring develop quicker under warmer
maternity roosting conditions [43]. Due to the ongoing pressure to be ready for migration and
subsequent hibernation, fast development is important for juvenile survivorship [44,45].
Therefore, a frequently appearing narrative is that reproductive females select roost in trees
located at lower elevations where conditions are warmer [40,46,47]. Our results point to dif-
ferent effect of elevation, that eastern red bats, especially reproductive females, roost at higher
elevations, but with the same inference: that female bats often chose to roost in warmer areas
on the landscape. Unlike the studies cited above, lower elevations were associated with lower
temperatures in our study area. This difference in the thermal gradient likely stems from the
scale of topography between our sites in southeastern Ohio, where elevation only varied by
approximately 100 m, often in the form of narrow, steep gorges. Despite the subtle difference
in elevation in our study, temperatures differed by ~3˚ C from ridges to bottoms, a biologically
meaningful difference from a thermoregulatory perspective.
It is notable that females at Crane Hollow Preserve were more likely to select roosts at
higher elevations in comparison to males. Sexual segregation in bats has been observed by oth-
ers, in which, females occupied lower, warmer elevations, than males during pregnancy and
lactation [40,46,47]. Males are less energetically constrained than females, and likely experi-
ence fewer costs associated with roosting at lower temperatures [17,48]. However, lesser ener-
getic constraints in males does not fully explain why males at Crane Hollow Preserve roosted
at a different range of elevations than females, as roosting at colder elevations would increase
the cost of maintaining normothermy for males. This finding suggests either that lower eleva-
tions provided males with a benefit such as the ability to use deeper torpor or the opportunity
to avoid competition with bats in higher quality (warmer) microclimates.
Meta-analyses and reviews of day-roost selection consistently find that tree diameter is a
primary predictor of whether a tree is used by bats [29,41]. Larger diameter trees are thought
to benefit bark and cavity roosting species because these trees provide more room for roosting
groups of bats [16,22]. Larger trees have also been hypothesized to have thicker bark and are
sometimes located in more open areas, making the tree a better thermal environment for
roosting [49]. These explanations are unlikely to explain why red bats preferred larger diame-
ter trees at the preserve, as increased space for social roosting would not benefit solitary red
bats, and roost trees were not located in open habitats. It is also difficult to explain why males
would roost in larger trees than females. Tree diameter at this site was not correlated with
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elevation, and we saw no evidence that larger trees were available at lower elevations, which
could have otherwise helped to explain this difference. Thus, while it is apparent that several
aspects of larger trees can benefit other bat species, it’s unclear why foliage roosting species
would prefer larger diameter trees.
Although our results do not provide insight into why tree diameter is important for red
bats, it is clear from our results that sexes selected roosts differently when larger diameter trees
were available. However, at Sunday Creek Coal, where we caught fewer red bats overall and
females were rare, we found no evidence of sexual segregation. We hypothesize that this differ-
ence was the result of forest conditions in each area. We propose that dense forests of relatively
small diameter trees, such as that at Sunday Creek Coal, are marginal roosting habitats for east-
ern red bats. In such areas, males and females either fail to segregate where they otherwise
would, or female populations are so low as to eliminate the need for segregation, although fur-
ther studies are needed in this area. Regardless, our results provide clear support of previous
studies showing that forest structure is an important aspect of day-roosting habitat for the east-
ern red bat in southeastern Ohio [26,29,31].
Finally, distance to water was a variable in our top model at Sunday Creek Coal. Proximity
to water is important for many bats, which often drink and forage in the area surrounding
their roosts upon evening emergence [15,50]. However, we observed the opposite trend, with
both male and female eastern red bats roosting farther from permanent water sources than
available trees. Because it is unlikely that bats seek to roost far from water, it is likely that this
metric either failed to capture water availability or was correlated with another, unmeasured
variable. Ephemeral water sources regularly used by bats, such as puddles in forest roads or
vernal pools, were found at higher elevations but were not be accounted for in our study.
Regardless, the average distance measured between the water sources and roosts was 200 m, a
distance easily traversed by bats, and unlikely to impact them negatively [51].
The underlying motivation for many studies of bat habitat selection is to guide manage-
ment practices. Over the last several decades, a large body of work has emerged describing the
habitat of different species in different regions, with some clear trends emerging [2,29,41].
Unfortunately, many of these trends are aggregated across species and are biased towards stud-
ies of cavity- and bark-roosting species. However, foliage roosting species are also experiencing
significant population declines, and more research is needed to guide management for these
species as well. While our netting data cannot be used to compare relative abundance of each
sex in our study areas, it is notable that we only captured 7 female red bats at Sunday Creek
Coal over four years of effort. It is therefore likely that females are rare in this area. Regional
based differences in sex ratios are common for bat species and have been well-documented in
eastern red bats. Although it is not uncommon for surveys to document these male-biased sex
ratios, data are often insufficient to make clear conclusions why [40,46,47]. Temperature as a
function of elevation [40,46,47], resource distribution [46], and seasonal patterns [52] have
often been the primary hypotheses explaining these differences. However, given the close
proximities of our study locations and the similarities in temperature regimes, habitat quality
may be another variable to consider when parsing out what is driving some of these patterns.
We encourage researchers to study habitat use within different forest contexts, beyond differ-
ent silvicultural treatments, and over different temperature regimes. Investigations of where
female reproduction does and does not occur in reference to these characteristics across a spe-
cies’ range can lead to better understanding of factors necessary for reproduction, driving hab-
itat use, and helping guide management and conservation efforts.
In the absence of these studies, our work highlights the importance of maintaining diversity
in forest structure over elevational gradients for eastern red bats in southeastern Ohio. We sug-
gest retaining trees >38 cm DBH (based on our overall average roost diameter at Crane
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Hollow Preserve) across an elevational gradient to provide suitable roost trees for eastern red
bats. To improve habitat in forests such as Sunday Creek Coal, single tree selective thinning or
group selection thinning could be employed to open the understory and release remaining
trees from competition. Despite our data suggesting that sugar maples were the only tree spe-
cies used more than expected by females, we also recommended retaining tulip poplars, oaks,
and sweet birch since these species were frequently used at both study sites, indicating they are
suitable roosts. While the specific effects of the variables we report must be viewed in the con-
text of our study areas where the topography created cooler temperatures in riparian areas and
at the bottom of gorges, the inference that temperature has a strong influence on habitat selec-
tion by eastern red bats may broadly apply throughout the species’ range. We urge land man-
agers to provide roosting habitat in relatively warm microclimates where thermal conditions
vary on the local conditions.
Supporting information
S1 Table. Biologically informed a priori models and corresponding variables used for our
information theoretic approach and multinomial logistic regressions. Superscripts provide
citations justifying the variables in each model.
(PNG)
Acknowledgments
We would like to thank H. Stehle and J. Moosbrugger from Crane Hollow, B. Brown, R. Har-
ris, E. Hazelton, J. Norris, and K. Parsons of the ODNR, and K. Schultes of the United States
Forest Service for their support. We would also like to thank B. Wittich, J. Geiger, E. Ebert, I.
Ruta, H. Slagel, T. Black, K. Kelly, L. Williams, A. Shields, B. Cordle, and B. Rechel for their
assistance in the field.
Author Contributions
Conceptualization: Joseph S. Johnson.
Data curation: Maria N. Monarchino, Marnie L. Behan, Joseph S. Johnson.
Formal analysis: Maria N. Monarchino.
Funding acquisition: Joseph S. Johnson.
Investigation: Maria N. Monarchino, Marnie L. Behan.
Methodology: Maria N. Monarchino, Marnie L. Behan, Joseph S. Johnson.
Project administration: Maria N. Monarchino, Joseph S. Johnson.
Visualization: Maria N. Monarchino, Marnie L. Behan.
Writing – original draft: Maria N. Monarchino, Joseph S. Johnson.
Writing – review & editing: Maria N. Monarchino, Marnie L. Behan, Joseph S. Johnson.
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