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Northern Long-Eared Bat (Myotis septentrionalis) Day-Roost Loss in the Central Appalachian Mountains following Prescribed Burning

Northern Long-Eared Bat (Myotis septentrionalis) Day-Roost Loss in the Central Appalachian... Hindawi International Journal of Forestry Research Volume 2021, Article ID 5512044, 6 pages https://doi.org/10.1155/2021/5512044 Research Article Northern Long-Eared Bat (Myotis septentrionalis) Day-Roost Loss in the Central Appalachian Mountains following Prescribed Burning 1 2 3 W. Mark Ford , Joshua B. Johnson, and Melissa Thomas-Van Gundy U.S. Geological Survey, Virginia Cooperative Fish and Wildlife Research Unit, Blacksburg, VA 24061, USA Pennsylvania Game Commission, Harrisburg, PA 17110, USA U.S. Forest Service, Northern Research Station, Parsons, WV 26287, USA Correspondence should be addressed to W. Mark Ford; wmford@vt.edu Received 1 March 2021; Revised 17 June 2021; Accepted 23 June 2021; Published 2 July 2021 Academic Editor: Nikolaos D. Hasanagas Copyright © 2021 W. Mark Ford et al. ,is is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Before the arrival of white-nose syndrome in North America, the northern long-eared bat (Myotis septentrionalis) was a common cavity-roosting bat species in central Appalachian hardwood forests. Two successive prescribed burns on the Fernow Experi- mental Forest, West Virginia, in 2008 and 2009, were shown to positively affect maternity colony day-roost availability and condition in the near-term. However, whether immediate benefits were temporary and if burned forests actually experienced an accelerated loss of trees and snags possibly suitable for bats more than background loss in unburned forests became an important question following the species’ threatened designation. In 2016, we revisited 81 of 113 northern long-eared bat maternity colony day-roosts initially discovered in 2007–2009 with the objective of ascertaining if these trees and snags were still standing and thus potentially “available” for bat use. Initial tree or snag stage condition class and original year of discovery were contributory factors determining availability by 2016, whereas exposure to prescribed fire and tree/snag species decay resistance were not. Because forest managers may consider using habitat enhancement to improve northern long-eared bat survival, reproduction, and juvenile recruitment and must also protect documented day-roosts during forestry operations, we conclude that initial positive benefits from prescribed burning did not come at the expense of subsequent day-roost loss greater than background rates in these forests at least for the duration we examined. suitable and often ephemeral conditions can be rare re- 1. Introduction gionally in maturing deciduous forests [5]. Return of fire to Conservation of day-roosts is a central management tenet the central Appalachian landscape through prescribing for many bat species. In much of eastern North America, burning has been shown to improve foraging habitat con- particularly the heavily forested central and southern Ap- ditions for bats generally by reducing forest “clutter” such as palachian Mountains, lack of forests and/or trees and snags midstory growth [6, 7]. is not a limiting factor per se as much as forest stand For day-roost conditions, burning improves, at least structure that provides optimal condition and spatial con- temporarily, conditions for Indiana bats and the threatened figuration of suitable roosts. For example, endangered northern long-eared bats (Myotis septentrionalis) [5, 8]. In Indiana bat (Myotis sodalis) summer maternity colonies the maternity season, female northern long-eared bat day- often require recently fire or flood-disturbed older-age roosts are in cavities of live trees or standing snags across a stands containing trees and snags with exfoliating bark and wide variation in bole size and solar exposure [9]. Although high solar exposure as their ideal roost conditions are some northern long-eared bat day-roosts are lost from ephemeral and transitory [1–4]. Unfortunately, with dis- collapse or combustion during a prescribed burn [8], re- rupted disturbance regimes, i.e., fire suppression, these peated fires can accelerate the processes of both cavity 2 International Journal of Forestry Research formation and transitioning a healthy tree to a declining tree from 530 m to 1100 m [19]. Mean annual precipitation at or snag [10] that is more suitable for a day-roost [11]. FEF is 145.8 cm and mean annual temperature is 9.2 C, with Similarly, in the near-term, northern long-eared bat ma- an average frost-free period of 145 days [20]. Forest stands ternity colonies select day-roosts in burned stands more than on the FEF are a mosaic of second- and third-growth, mixed- expected relative to unburned stands based on availability mesophytic, and northern hardwood types that have been [8]. Ford et al. [12] observed a postfire reduction in trees and managed by various even- and uneven-aged harvesting standing snags of species favored by northern long-eared practices or have been left undisturbed following initial bats. However, the proportion of trees and snags in advanced harvesting in the early 20th century [19, 21]. On parts of the stages of decline or decay suitable for northern long-eared FEF and much of the surrounding Monongahela National bat day-roosts increased postfire among highly used tree and Forest, American chestnut (Castanea dentata) and oaks snag species such as black locust (Robinia pseudoacacia) (Quercus spp.), such as northern red oak (Quercus rubra), relative to unburned stands. Additionally, maternity colony historically dominated the forest overstory. However, social cohesion, as measured by the centrality of day-roosts chestnut blight (Cryphonectria parasitica) and subsequent used and numbers shared by conspecifics, was greater in lack of intense disturbance, including fire and harvesting, burned stands than unburned stands [12]. since the mid-20th century has allowed forest composition ,e use of fire as a forest management tool in the central in unmanaged stands to shift toward shade-tolerant tree Appalachians typically is targeted towards other stewardship species, such as maples (Acer spp.) and American beech needs such as controlling shade-tolerant competition to (Fagus grandifolia) [21]. Since 2002, prescribed fire has been foster oak (Quercus spp.) regeneration or to maintain early used to promote oak regeneration in some FEF forest stands successional habitat types [13, 14]. Although evidence where the regeneration layer is currently dominated by sugar suggests that burning can provide benefits to bats in the maple (Acer saccharum), red maple (Acer rubrum), yellow- region, questions remain about the longevity of bat day- poplar (Liriodendron tulipifera), black cherry (Prunus roosts and the long-term effects following fire [15]. Dis- serotina), American beech, sweet birch (Betula lenta), and ruption of northern long-eared bat maternity colony social basswood (Tilia americana) [8, 22]. In early spring of 2007 networks can be caused by day-roost losses exceeding 30% and 2008, consecutive prescribed fires were conducted in a from one year to the next [16]. Because this species has been 121 ha compartment on the FEF that had been relatively free heavily impacted by white-nose syndrome, with population of management except for scattered light timber stand declines exceeding 90% in the central Appalachians [17], improvement entries in the previous decades. Treatment forest managers now seek to enhance summer maternity stands were burned using a strip head fire technique, ignited day-roosting habitat for this species to promote successful with handheld drip torches after fire-blackened perimeters reproduction and juvenile recruitment [18]. For northern were established [12]. long-eared bats, it is not known if prescribed fire provides only short-term day-roost benefits followed by an 2.2. Data Collection. During the summers of 2007–2009, we accelerated loss relative to background conditions. ,is could have conservation implications for the use of fire in mist-netted and captured northern long-eared bats on the forests where northern long-eared bats occur or the use of FEF in and around the burned compartment. Captured adult females were outfitted with radio-transmitters and tracked fire as a specific bat habitat management tool. Accordingly, in 2016, we tracked fate (standing or fallen) to their maternity colony day-roosts for the life of the transmitter or until the transmitter fell off. For complete of northern long-eared bat maternity colony day-roost trees and snags recorded from 2007 to 2009 at the Fernow Ex- details on mist-netting and tracking techniques, see [8, 11, 12]. Numerous metrics were recorded to describe and perimental Forest (FEF) in northcentral West Virginia [8, 11, 12] in burned and unburned areas. We predicted that differentiate day-roost characteristics between the burned compartment and adjacent unburned stands such as di- the probability of a day-roost tree or snag remaining in 2016 would be negatively related to initial tree or snag condition ameter at breast height, crown class, condition class, bark retention, cavity presence, roost type (cavity or bark), and class when first discovered, positively related to tree or snag species wood specific gravity as a surrogate for wood decay tree height of day-roost as well as surrounding basal area, resistance, negatively related to exposure to fire, and neg- and these data are presented elsewhere [8, 11, 12]. Because all located day-roosts from 2007 to 2009 were georeferenced atively related to time of original discovery. and tagged, in the summer of 2016, U.S. Forest Service, Northern Research Station staff relocated a large portion of 2. Methods the day-roosts to establish necessary protective buffer zones pursuant to the requirements in the northern long-eared bat 2.1. Study Area. We examined data for northern long-eared bat day-roosts that were originally collected by [5, 8] from 4(d) rule [23]. Located day-roosts were then recorded as live tree, standing snag, or fallen. No other measurements were the FEF in northeastern West Virginia (39 03′15″N 79 41′15″W). ,e FEF is a 1,900 ha experimental forest collected nor were day-roosts checked for continued oc- cupancy by northern long-eared bats or any other myotids, managed by the U.S. Forest Service, Northern Research Station, located in the unglaciated Allegheny Plateau portion as the impacts of WNS and subsequent precipitous pop- ulation declines ∼90% observed in Big Springs Cave, a of the central Appalachians. Topography is characterized by sizable bat hibernaculum on the FEF, made that unlikely. It steep slopes and plateau-like ridgetops. Elevations range International Journal of Forestry Research 3 is possible that the other cavity-roosting bat species present, the top supported model, there was no difference in standing the big brown bat (Eptesicus fuscus), used these as day- versus fallen outcomes attributable to exposure to prescribed roosts; however, none were noted roosting in earlier surveys burning (Table 2, Figure 1). Both original year of day-roost [8], and their presence on FEF is relatively low [24]. discovery and initial condition class affected day-roost availability by 2016, though parameter estimate confidence intervals just crossed zero (Table 2). Day-roosts discovered 2.3. Data Analysis. To assess the effect of being subjected to initially in 2009 were less likely to remain standing in 2016 prescribed fire or unburned (FIRE), initial tree or snag than those from 2007 to 2008 (Figure 1). Day-roosts in condition class (STAGE: 1–7) [25], surrogate decay resis- higher condition classes, i.e., live trees in decay stage 2 or tance rank from tree species specific gravity values collected snags with a higher degree of decay, upon initial discovery at the FEF (RANK) [26], diameter at breast height (DBH), 2007–2009, were more likely to have fallen by 2016 and year of initial discovery (YEAR: 2007–2009), we (Figure 1). modeled the outcome of day-roosts remaining standing and presumably available versus fallen. Ideally, these data would 4. Discussion be analyzed using a survival analysis approach [27]; how- ever, without annual assessments from 2009 to 2016, this was Ford et al. [8, 12] showed that relatively small-sized live trees not possible. ,erefore, because we could only assess the and trees or snags in decay condition classes 2–6 [25] final time step outcome, we used a generalized linear model composed approximately 50% of day-roosts used on the FEF with a binomial distribution and a logit link function (PROC by northern long-eared bats. Black locust along with red GENMOD, SAS 10.2, Cary, NC). Surrogate decay resistance maple, northern red oak, and black oak were the only species ranks were different among all species of trees and snags; selected more than availability would suggest from the FEF’s therefore, this variable also served as a surrogate for species. diverse tree species pool [5, 8, 12]. Irrespective of the in- Because determining if fire had an effect on day-roost troduction of fire, abundant numbers of small to medium- availability was our primary question, all candidate models sized black locust and sassafras trees and snags with cavities contained the FIRE variable. With FIRE, we modeled and suitable for northern long-eared bat day-roosts would be ranked all combinations of the four variables (STAGE, common in these FEF stands owing to the stand age and RANK, DBH, and YEAR) using Akaike’s information cri- known patterns of stand development in the central Ap- teria (AIC) value corrected for small sample size [28]. palachians [29]. Although some day-roosts located initially in 2007 were snags that predated the two prescribed burns on the FEF, northern long-eared bats in 2008 and 2009 did 3. Results use snags that were a direct result of tree mortality following Of 113 northern long-eared bat day-roosts recorded from fire [8, 12]. 2007 to 2009, 81 were relocated in the 2016 search effort with A preponderance of research, mostly from western 78 being used in our analyses. ,ree of these day-roosts had North America, suggests that longevity of trees that become been cut from long-term silvicultural studies prior to the snags or current snags following fire can be one-third to one- listing of the northern long-eared bat and were excluded. Of half as long as that in unburned stands [30]. ,ese com- the 15 that were originally live trees, 12 remained alive and parisons are relatively limited for deciduous forests where we worked or conifer, i.e., southern pine (Pinus spp.) or standing in 2016, 2 had become standing dead snags, and 1 tree had fallen. Of the 63 that were originally standing snags, boreal systems in the East. Our data show no substantive 39 remained standing in 2016, whereas 24 had fallen. Trees difference in the probability of a burned tree or snag and snags that remained available as day-roosts averaged remaining available as a day-roost over time as compared to 29.8± 9.8 cm diameter at breast height at the time of original unburned trees at the FEF. In hardwood forests, surprisingly, discovery, whereas those that had fallen averaged snag persistence across most size classes of boles and tree 26.6± 6.8 cm. Black locust, red maple, and sassafras (Sas- species may only be about 10 years in the southern Piedmont safras albidum) comprised approximately 70% of the day- [31] and 20–25 years in the northern Appalachians [32]. At roosts relocated in 2016 with 40%, 36%, and 38%, respec- the FEF, either the progression from tree to decaying snag to tively, becoming unavailable by 2016. Conversely, of the 10 fallen coarse woody debris may occur quickly; masking fire northern red oaks, 3 chestnut oaks (Quercus prinus), one effects on our resampling effort in 2016 had not allowed enough time to elapse for differentiation among rates of black oak (Quercus velutina), and one white oak (Quercus alba) monitored through 2016, only a single northern red senescence. Despite a slightly higher probability of roost oak, originally in decay stage 3, and the one white oak, trees remaining available overall in unburned stands, the originally in decay stage 6, had fallen by 2016. rates of change of availability between unburned and burned ,e model retaining fire, initial decay stage, and year of in our study were similar. For the most part, the loss of day- initial day-roost discovery was the top supported model roosts following the two successive prescribed fires on the (Table 1). Day-roost species or size were not as important, as FEF probably did not cause loss more than background rates no model containing decay resistance rank or diameter at that were normally occurring as part of ongoing forest breast height was within ΔAIC < 2. ,e only other variable dynamic processes. Year of initial discovery was a factor combinations with support were fire and year of initial day- contributing to long-term availability of day-roosts. ,ose roost discovery or fire and initial decay stage (Table 1). From initially found in 2009 were less likely to be available to 4 International Journal of Forestry Research Table 1: Competing generalized linear models and associated number of parameters (K), Akaike’s information criterion values (AIC ), model rankings (ΔAIC ), and Akaike weights (ω ) for predicting availability of northern long-eared bat (Myotis septentrionalis) day-roosts c i discovered in 2007–2009 and reexamined in 2016 on the Fernow Experimental Forest (FEF), West Virginia. Model K AIC ΔAIC ω c c i FIRE + STAGE + YEAR 4 97.81 0.00 0.25 FIRE + YEAR 3 99.16 1.35 0.13 FIRE + STAGE 3 99.17 1.36 0.13 FIRE + DBH + RANK + YEAR 5 99.96 2.15 0.09 FIRE 2 100.03 2.22 0.08 FIRE + RANK + STAGE + YEAR 5 100.07 2.26 0.08 FIRE + DBH + STAGE + YEAR 5 100.64 2.83 0.06 FIRE + DBH + STAGE 4 101.22 3.41 0.05 FIRE + DBH + YEAR 4 101.27 3.46 0.04 FIRE + RANK + STAGE 4 101.35 3.54 0.04 FIRE + RANK + YEAR 4 101.39 3.58 0.04 FIRE + DBH 3 101.98 4.17 0.03 FIRE + RANK 3 102.23 4.42 0.03 GLOBAL (FIRE + DBH + RANK + STAGE + YEAR) 6 102.28 4.47 0.03 FIRE + DBH + RANK + STAGE 5 103.52 5.71 0.01 FIRE + DBH + RANK 4 104.09 6.28 0.01 FIRE, prescribed fire vs. unburned; STAGE, initial day-roost decay class condition; YEAR, year of day-roost discovery; RANK, surrogate decay resistance rank from tree species specific gravity values collected at the FEF; DBH, diameter at breast height of day-roost at initial discovery. Table 2: Parameter estimates of the best approximating model for predicting availability of northern long-eared bat (Myotis septentrionalis) day-roosts discovered in 2007–2009 and reexamined in 2016 on the Fernow Experimental Forest, West Virginia. Wald 95% confidence a 2 Variable d.f. Parameter estimate Standard error Wald χ P value limits Intercept 1 1765.80 959.22 −114.24 3645.84 3.39 0.07 Fire 1 0.48 0.54 −0.58 1.54 0.79 0.37 Stage 1 −0.32 0.18 −0.67 0.02 3.33 0.07 Year 1 −0.88 0.80 −1.81 0.06 3.38 0.07 FIRE, prescribed fire vs. unburned; STAGE, initial day-roost decay class condition; YEAR, year of day-roost discovery; RANK, surrogate decay resistance rank from tree species specific gravity values collected at the FEF; DBH, diameter at breast height of day-roost at initial discovery. 1.0 0.8 Unburned 0.6 Burned 0.4 0.2 0.0 1234 56 7 Tree/snag starting decay stage 1–7 Figure 1: Probability of northern long-eared bat (Myotis septentrionalis) day-roosts being available (standing) in 2016 relative to exposure to two prescribed fires and initial day-roost decay stage on the Fernow Experimental Forest, West Virginia, 2007–2009. Shading represents the 95% credible estimate around the prediction. northern long-eared bats by 2016. However, we attribute this without being documented, as northern long-eared bat paradox to the relatively small number (4) of new day-roosts, maternity colonies often will occupy the same relatively only in burned stands, added in 2009. It is possible, of course, small general location for successive years in temperate that these day-roosts were also used in 2007 and 2008 deciduous forests so long as day-roosts are suitable [9]. Probability of availability International Journal of Forestry Research 5 What remains unknown from our assessment of day- Disclosure roost availability on the FEF is the annual to long-term trend Any use of trade, firm, or product names is for descriptive in recruitment and loss in live trees and snags with cavities purposes only and does not imply endorsement by the U.S. suitable for northern long-eared bats. Cavity tree and snag Government. abundance can be highly variable in the central and southern Appalachians depending on forest type [33], though often, there is no relationship between these and forest stand age Conflicts of Interest [34]. Archived data from an unmanaged watershed on the ,e authors declare that they have no conflicts of interest. FEF showed there was an annual recruitment of 0.4 snags/ ha/year >12.7 cm. In mid-1960s, there was an average of 15.5 snags/ha increasing to 38.1/ha by 2019, densities greater than Authors’ Contributions per area needs of northern long-eared bat maternity colonies WMF and JBJ conceived the study. WMF provided study [35]. Subtle changes in snag species are apparent over these oversight, performed the analyses, and prepared the man- 50+ years with early successional black locust and sassafras uscript. JBJ and MTV collected field data. WMF, JBJ, and comprising a large proportion of snags initially and then oak MTV interpreted the data and edited and prepared the snags increasing in three decades later, a group highly se- manuscript for submission. lected as day-roosts for the northern long-eared bat in the southern Appalachians [36]. Acknowledgments 5. Conclusions ,e authors thank D. Lowther, R. Hovatter, M. Karp, J. Rodrigue, T. Schuler, and numerous FEF summer interns Some forest dwelling bat species require a shifting mosaic for field assistance throughout this effort. Financial support across the landscape of some ephemeral trees or stand for this project was provided by U.S. Forest Service, conditions [5, 37]. At the landscape scale in central Appa- Northern Research Station. lachian oak-pine forests, prescribed fire has been shown capable of providing that stand condition heterogeneity [38]. 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[19] W. M. Ford, M. A. Menzel, J. L. Rodrigue, J. M. Menzel, and [36] J. M. O’Keefe, Roosting and foraging ecology of forest bats in J. B. Johnson, “Relating bat species presence to simple habitat the southern Appalachian Mountains, Ph.D. Dissertation, measures in a central Appalachian forest,” Biological Con- Clemson University, Clemson, SC, USA, 2009. servation, vol. 126, no. 4, pp. 528–539, 2005. [37] K. J. Bondo, C. K. R. Willis, J. D. Metheny et al., “Bats relocate maternity colony after the natural loss of roost trees,” 7e [20] D. M. Madarish, J. L. Rodrigue, and M. B. Adams, “Vascular flora and macroscopic fauna on the Fernow experimental Journal of Wildlife Management, vol. 83, no. 8, pp. 1753–1761, forest,” Technical Report NE-291, USDA Forest Service, 2019. Northeastern Research Station, Newtown Square, PA, USA, [38] J. Lorber, M. ,omas-Van Gundy, and S. Croy, Characterizing 2002. Effects of Prescribed Fire on Forest Canopy Cover in the George [21] T. M. Schuler, “Fifty years of partial harvesting in a mixed Washington and Jefferson National Forests, USDA Forest mesophytic forest: composition and productivity,” Canadian Service Northern Research Station Research Paper. NRS-RP- Journal of Forest Research, vol. 34, no. 5, pp. 985–997, 2004. 31, Newtown Square, PA, USA, 2018. [22] W. M. Ford, S. F. Owen, J. W. Edwards, and J. L. Rodrigue, [39] M. S. Muthersbaugh, “Seasonal activity patterns of bats in the “Robinia pseudoacacia (black locust) as day-roosts of male central appalachians,” M. S. thesis, Virginia Polytechnic In- Myotis septentrionalis (northern bats) on the Fernow stitute and State University, Blacksburg, VA, USA, 2018. http://www.deepdyve.com/assets/images/DeepDyve-Logo-lg.png International Journal of Forestry Research Hindawi Publishing Corporation

Northern Long-Eared Bat (Myotis septentrionalis) Day-Roost Loss in the Central Appalachian Mountains following Prescribed Burning

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Copyright © 2021 W. Mark Ford et al. This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
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Hindawi International Journal of Forestry Research Volume 2021, Article ID 5512044, 6 pages https://doi.org/10.1155/2021/5512044 Research Article Northern Long-Eared Bat (Myotis septentrionalis) Day-Roost Loss in the Central Appalachian Mountains following Prescribed Burning 1 2 3 W. Mark Ford , Joshua B. Johnson, and Melissa Thomas-Van Gundy U.S. Geological Survey, Virginia Cooperative Fish and Wildlife Research Unit, Blacksburg, VA 24061, USA Pennsylvania Game Commission, Harrisburg, PA 17110, USA U.S. Forest Service, Northern Research Station, Parsons, WV 26287, USA Correspondence should be addressed to W. Mark Ford; wmford@vt.edu Received 1 March 2021; Revised 17 June 2021; Accepted 23 June 2021; Published 2 July 2021 Academic Editor: Nikolaos D. Hasanagas Copyright © 2021 W. Mark Ford et al. ,is is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Before the arrival of white-nose syndrome in North America, the northern long-eared bat (Myotis septentrionalis) was a common cavity-roosting bat species in central Appalachian hardwood forests. Two successive prescribed burns on the Fernow Experi- mental Forest, West Virginia, in 2008 and 2009, were shown to positively affect maternity colony day-roost availability and condition in the near-term. However, whether immediate benefits were temporary and if burned forests actually experienced an accelerated loss of trees and snags possibly suitable for bats more than background loss in unburned forests became an important question following the species’ threatened designation. In 2016, we revisited 81 of 113 northern long-eared bat maternity colony day-roosts initially discovered in 2007–2009 with the objective of ascertaining if these trees and snags were still standing and thus potentially “available” for bat use. Initial tree or snag stage condition class and original year of discovery were contributory factors determining availability by 2016, whereas exposure to prescribed fire and tree/snag species decay resistance were not. Because forest managers may consider using habitat enhancement to improve northern long-eared bat survival, reproduction, and juvenile recruitment and must also protect documented day-roosts during forestry operations, we conclude that initial positive benefits from prescribed burning did not come at the expense of subsequent day-roost loss greater than background rates in these forests at least for the duration we examined. suitable and often ephemeral conditions can be rare re- 1. Introduction gionally in maturing deciduous forests [5]. Return of fire to Conservation of day-roosts is a central management tenet the central Appalachian landscape through prescribing for many bat species. In much of eastern North America, burning has been shown to improve foraging habitat con- particularly the heavily forested central and southern Ap- ditions for bats generally by reducing forest “clutter” such as palachian Mountains, lack of forests and/or trees and snags midstory growth [6, 7]. is not a limiting factor per se as much as forest stand For day-roost conditions, burning improves, at least structure that provides optimal condition and spatial con- temporarily, conditions for Indiana bats and the threatened figuration of suitable roosts. For example, endangered northern long-eared bats (Myotis septentrionalis) [5, 8]. In Indiana bat (Myotis sodalis) summer maternity colonies the maternity season, female northern long-eared bat day- often require recently fire or flood-disturbed older-age roosts are in cavities of live trees or standing snags across a stands containing trees and snags with exfoliating bark and wide variation in bole size and solar exposure [9]. Although high solar exposure as their ideal roost conditions are some northern long-eared bat day-roosts are lost from ephemeral and transitory [1–4]. Unfortunately, with dis- collapse or combustion during a prescribed burn [8], re- rupted disturbance regimes, i.e., fire suppression, these peated fires can accelerate the processes of both cavity 2 International Journal of Forestry Research formation and transitioning a healthy tree to a declining tree from 530 m to 1100 m [19]. Mean annual precipitation at or snag [10] that is more suitable for a day-roost [11]. FEF is 145.8 cm and mean annual temperature is 9.2 C, with Similarly, in the near-term, northern long-eared bat ma- an average frost-free period of 145 days [20]. Forest stands ternity colonies select day-roosts in burned stands more than on the FEF are a mosaic of second- and third-growth, mixed- expected relative to unburned stands based on availability mesophytic, and northern hardwood types that have been [8]. Ford et al. [12] observed a postfire reduction in trees and managed by various even- and uneven-aged harvesting standing snags of species favored by northern long-eared practices or have been left undisturbed following initial bats. However, the proportion of trees and snags in advanced harvesting in the early 20th century [19, 21]. On parts of the stages of decline or decay suitable for northern long-eared FEF and much of the surrounding Monongahela National bat day-roosts increased postfire among highly used tree and Forest, American chestnut (Castanea dentata) and oaks snag species such as black locust (Robinia pseudoacacia) (Quercus spp.), such as northern red oak (Quercus rubra), relative to unburned stands. Additionally, maternity colony historically dominated the forest overstory. However, social cohesion, as measured by the centrality of day-roosts chestnut blight (Cryphonectria parasitica) and subsequent used and numbers shared by conspecifics, was greater in lack of intense disturbance, including fire and harvesting, burned stands than unburned stands [12]. since the mid-20th century has allowed forest composition ,e use of fire as a forest management tool in the central in unmanaged stands to shift toward shade-tolerant tree Appalachians typically is targeted towards other stewardship species, such as maples (Acer spp.) and American beech needs such as controlling shade-tolerant competition to (Fagus grandifolia) [21]. Since 2002, prescribed fire has been foster oak (Quercus spp.) regeneration or to maintain early used to promote oak regeneration in some FEF forest stands successional habitat types [13, 14]. Although evidence where the regeneration layer is currently dominated by sugar suggests that burning can provide benefits to bats in the maple (Acer saccharum), red maple (Acer rubrum), yellow- region, questions remain about the longevity of bat day- poplar (Liriodendron tulipifera), black cherry (Prunus roosts and the long-term effects following fire [15]. Dis- serotina), American beech, sweet birch (Betula lenta), and ruption of northern long-eared bat maternity colony social basswood (Tilia americana) [8, 22]. In early spring of 2007 networks can be caused by day-roost losses exceeding 30% and 2008, consecutive prescribed fires were conducted in a from one year to the next [16]. Because this species has been 121 ha compartment on the FEF that had been relatively free heavily impacted by white-nose syndrome, with population of management except for scattered light timber stand declines exceeding 90% in the central Appalachians [17], improvement entries in the previous decades. Treatment forest managers now seek to enhance summer maternity stands were burned using a strip head fire technique, ignited day-roosting habitat for this species to promote successful with handheld drip torches after fire-blackened perimeters reproduction and juvenile recruitment [18]. For northern were established [12]. long-eared bats, it is not known if prescribed fire provides only short-term day-roost benefits followed by an 2.2. Data Collection. During the summers of 2007–2009, we accelerated loss relative to background conditions. ,is could have conservation implications for the use of fire in mist-netted and captured northern long-eared bats on the forests where northern long-eared bats occur or the use of FEF in and around the burned compartment. Captured adult females were outfitted with radio-transmitters and tracked fire as a specific bat habitat management tool. Accordingly, in 2016, we tracked fate (standing or fallen) to their maternity colony day-roosts for the life of the transmitter or until the transmitter fell off. For complete of northern long-eared bat maternity colony day-roost trees and snags recorded from 2007 to 2009 at the Fernow Ex- details on mist-netting and tracking techniques, see [8, 11, 12]. Numerous metrics were recorded to describe and perimental Forest (FEF) in northcentral West Virginia [8, 11, 12] in burned and unburned areas. We predicted that differentiate day-roost characteristics between the burned compartment and adjacent unburned stands such as di- the probability of a day-roost tree or snag remaining in 2016 would be negatively related to initial tree or snag condition ameter at breast height, crown class, condition class, bark retention, cavity presence, roost type (cavity or bark), and class when first discovered, positively related to tree or snag species wood specific gravity as a surrogate for wood decay tree height of day-roost as well as surrounding basal area, resistance, negatively related to exposure to fire, and neg- and these data are presented elsewhere [8, 11, 12]. Because all located day-roosts from 2007 to 2009 were georeferenced atively related to time of original discovery. and tagged, in the summer of 2016, U.S. Forest Service, Northern Research Station staff relocated a large portion of 2. Methods the day-roosts to establish necessary protective buffer zones pursuant to the requirements in the northern long-eared bat 2.1. Study Area. We examined data for northern long-eared bat day-roosts that were originally collected by [5, 8] from 4(d) rule [23]. Located day-roosts were then recorded as live tree, standing snag, or fallen. No other measurements were the FEF in northeastern West Virginia (39 03′15″N 79 41′15″W). ,e FEF is a 1,900 ha experimental forest collected nor were day-roosts checked for continued oc- cupancy by northern long-eared bats or any other myotids, managed by the U.S. Forest Service, Northern Research Station, located in the unglaciated Allegheny Plateau portion as the impacts of WNS and subsequent precipitous pop- ulation declines ∼90% observed in Big Springs Cave, a of the central Appalachians. Topography is characterized by sizable bat hibernaculum on the FEF, made that unlikely. It steep slopes and plateau-like ridgetops. Elevations range International Journal of Forestry Research 3 is possible that the other cavity-roosting bat species present, the top supported model, there was no difference in standing the big brown bat (Eptesicus fuscus), used these as day- versus fallen outcomes attributable to exposure to prescribed roosts; however, none were noted roosting in earlier surveys burning (Table 2, Figure 1). Both original year of day-roost [8], and their presence on FEF is relatively low [24]. discovery and initial condition class affected day-roost availability by 2016, though parameter estimate confidence intervals just crossed zero (Table 2). Day-roosts discovered 2.3. Data Analysis. To assess the effect of being subjected to initially in 2009 were less likely to remain standing in 2016 prescribed fire or unburned (FIRE), initial tree or snag than those from 2007 to 2008 (Figure 1). Day-roosts in condition class (STAGE: 1–7) [25], surrogate decay resis- higher condition classes, i.e., live trees in decay stage 2 or tance rank from tree species specific gravity values collected snags with a higher degree of decay, upon initial discovery at the FEF (RANK) [26], diameter at breast height (DBH), 2007–2009, were more likely to have fallen by 2016 and year of initial discovery (YEAR: 2007–2009), we (Figure 1). modeled the outcome of day-roosts remaining standing and presumably available versus fallen. Ideally, these data would 4. Discussion be analyzed using a survival analysis approach [27]; how- ever, without annual assessments from 2009 to 2016, this was Ford et al. [8, 12] showed that relatively small-sized live trees not possible. ,erefore, because we could only assess the and trees or snags in decay condition classes 2–6 [25] final time step outcome, we used a generalized linear model composed approximately 50% of day-roosts used on the FEF with a binomial distribution and a logit link function (PROC by northern long-eared bats. Black locust along with red GENMOD, SAS 10.2, Cary, NC). Surrogate decay resistance maple, northern red oak, and black oak were the only species ranks were different among all species of trees and snags; selected more than availability would suggest from the FEF’s therefore, this variable also served as a surrogate for species. diverse tree species pool [5, 8, 12]. Irrespective of the in- Because determining if fire had an effect on day-roost troduction of fire, abundant numbers of small to medium- availability was our primary question, all candidate models sized black locust and sassafras trees and snags with cavities contained the FIRE variable. With FIRE, we modeled and suitable for northern long-eared bat day-roosts would be ranked all combinations of the four variables (STAGE, common in these FEF stands owing to the stand age and RANK, DBH, and YEAR) using Akaike’s information cri- known patterns of stand development in the central Ap- teria (AIC) value corrected for small sample size [28]. palachians [29]. Although some day-roosts located initially in 2007 were snags that predated the two prescribed burns on the FEF, northern long-eared bats in 2008 and 2009 did 3. Results use snags that were a direct result of tree mortality following Of 113 northern long-eared bat day-roosts recorded from fire [8, 12]. 2007 to 2009, 81 were relocated in the 2016 search effort with A preponderance of research, mostly from western 78 being used in our analyses. ,ree of these day-roosts had North America, suggests that longevity of trees that become been cut from long-term silvicultural studies prior to the snags or current snags following fire can be one-third to one- listing of the northern long-eared bat and were excluded. Of half as long as that in unburned stands [30]. ,ese com- the 15 that were originally live trees, 12 remained alive and parisons are relatively limited for deciduous forests where we worked or conifer, i.e., southern pine (Pinus spp.) or standing in 2016, 2 had become standing dead snags, and 1 tree had fallen. Of the 63 that were originally standing snags, boreal systems in the East. Our data show no substantive 39 remained standing in 2016, whereas 24 had fallen. Trees difference in the probability of a burned tree or snag and snags that remained available as day-roosts averaged remaining available as a day-roost over time as compared to 29.8± 9.8 cm diameter at breast height at the time of original unburned trees at the FEF. In hardwood forests, surprisingly, discovery, whereas those that had fallen averaged snag persistence across most size classes of boles and tree 26.6± 6.8 cm. Black locust, red maple, and sassafras (Sas- species may only be about 10 years in the southern Piedmont safras albidum) comprised approximately 70% of the day- [31] and 20–25 years in the northern Appalachians [32]. At roosts relocated in 2016 with 40%, 36%, and 38%, respec- the FEF, either the progression from tree to decaying snag to tively, becoming unavailable by 2016. Conversely, of the 10 fallen coarse woody debris may occur quickly; masking fire northern red oaks, 3 chestnut oaks (Quercus prinus), one effects on our resampling effort in 2016 had not allowed enough time to elapse for differentiation among rates of black oak (Quercus velutina), and one white oak (Quercus alba) monitored through 2016, only a single northern red senescence. Despite a slightly higher probability of roost oak, originally in decay stage 3, and the one white oak, trees remaining available overall in unburned stands, the originally in decay stage 6, had fallen by 2016. rates of change of availability between unburned and burned ,e model retaining fire, initial decay stage, and year of in our study were similar. For the most part, the loss of day- initial day-roost discovery was the top supported model roosts following the two successive prescribed fires on the (Table 1). Day-roost species or size were not as important, as FEF probably did not cause loss more than background rates no model containing decay resistance rank or diameter at that were normally occurring as part of ongoing forest breast height was within ΔAIC < 2. ,e only other variable dynamic processes. Year of initial discovery was a factor combinations with support were fire and year of initial day- contributing to long-term availability of day-roosts. ,ose roost discovery or fire and initial decay stage (Table 1). From initially found in 2009 were less likely to be available to 4 International Journal of Forestry Research Table 1: Competing generalized linear models and associated number of parameters (K), Akaike’s information criterion values (AIC ), model rankings (ΔAIC ), and Akaike weights (ω ) for predicting availability of northern long-eared bat (Myotis septentrionalis) day-roosts c i discovered in 2007–2009 and reexamined in 2016 on the Fernow Experimental Forest (FEF), West Virginia. Model K AIC ΔAIC ω c c i FIRE + STAGE + YEAR 4 97.81 0.00 0.25 FIRE + YEAR 3 99.16 1.35 0.13 FIRE + STAGE 3 99.17 1.36 0.13 FIRE + DBH + RANK + YEAR 5 99.96 2.15 0.09 FIRE 2 100.03 2.22 0.08 FIRE + RANK + STAGE + YEAR 5 100.07 2.26 0.08 FIRE + DBH + STAGE + YEAR 5 100.64 2.83 0.06 FIRE + DBH + STAGE 4 101.22 3.41 0.05 FIRE + DBH + YEAR 4 101.27 3.46 0.04 FIRE + RANK + STAGE 4 101.35 3.54 0.04 FIRE + RANK + YEAR 4 101.39 3.58 0.04 FIRE + DBH 3 101.98 4.17 0.03 FIRE + RANK 3 102.23 4.42 0.03 GLOBAL (FIRE + DBH + RANK + STAGE + YEAR) 6 102.28 4.47 0.03 FIRE + DBH + RANK + STAGE 5 103.52 5.71 0.01 FIRE + DBH + RANK 4 104.09 6.28 0.01 FIRE, prescribed fire vs. unburned; STAGE, initial day-roost decay class condition; YEAR, year of day-roost discovery; RANK, surrogate decay resistance rank from tree species specific gravity values collected at the FEF; DBH, diameter at breast height of day-roost at initial discovery. Table 2: Parameter estimates of the best approximating model for predicting availability of northern long-eared bat (Myotis septentrionalis) day-roosts discovered in 2007–2009 and reexamined in 2016 on the Fernow Experimental Forest, West Virginia. Wald 95% confidence a 2 Variable d.f. Parameter estimate Standard error Wald χ P value limits Intercept 1 1765.80 959.22 −114.24 3645.84 3.39 0.07 Fire 1 0.48 0.54 −0.58 1.54 0.79 0.37 Stage 1 −0.32 0.18 −0.67 0.02 3.33 0.07 Year 1 −0.88 0.80 −1.81 0.06 3.38 0.07 FIRE, prescribed fire vs. unburned; STAGE, initial day-roost decay class condition; YEAR, year of day-roost discovery; RANK, surrogate decay resistance rank from tree species specific gravity values collected at the FEF; DBH, diameter at breast height of day-roost at initial discovery. 1.0 0.8 Unburned 0.6 Burned 0.4 0.2 0.0 1234 56 7 Tree/snag starting decay stage 1–7 Figure 1: Probability of northern long-eared bat (Myotis septentrionalis) day-roosts being available (standing) in 2016 relative to exposure to two prescribed fires and initial day-roost decay stage on the Fernow Experimental Forest, West Virginia, 2007–2009. Shading represents the 95% credible estimate around the prediction. northern long-eared bats by 2016. However, we attribute this without being documented, as northern long-eared bat paradox to the relatively small number (4) of new day-roosts, maternity colonies often will occupy the same relatively only in burned stands, added in 2009. It is possible, of course, small general location for successive years in temperate that these day-roosts were also used in 2007 and 2008 deciduous forests so long as day-roosts are suitable [9]. Probability of availability International Journal of Forestry Research 5 What remains unknown from our assessment of day- Disclosure roost availability on the FEF is the annual to long-term trend Any use of trade, firm, or product names is for descriptive in recruitment and loss in live trees and snags with cavities purposes only and does not imply endorsement by the U.S. suitable for northern long-eared bats. Cavity tree and snag Government. abundance can be highly variable in the central and southern Appalachians depending on forest type [33], though often, there is no relationship between these and forest stand age Conflicts of Interest [34]. Archived data from an unmanaged watershed on the ,e authors declare that they have no conflicts of interest. FEF showed there was an annual recruitment of 0.4 snags/ ha/year >12.7 cm. In mid-1960s, there was an average of 15.5 snags/ha increasing to 38.1/ha by 2019, densities greater than Authors’ Contributions per area needs of northern long-eared bat maternity colonies WMF and JBJ conceived the study. WMF provided study [35]. Subtle changes in snag species are apparent over these oversight, performed the analyses, and prepared the man- 50+ years with early successional black locust and sassafras uscript. JBJ and MTV collected field data. WMF, JBJ, and comprising a large proportion of snags initially and then oak MTV interpreted the data and edited and prepared the snags increasing in three decades later, a group highly se- manuscript for submission. lected as day-roosts for the northern long-eared bat in the southern Appalachians [36]. Acknowledgments 5. Conclusions ,e authors thank D. Lowther, R. Hovatter, M. Karp, J. Rodrigue, T. Schuler, and numerous FEF summer interns Some forest dwelling bat species require a shifting mosaic for field assistance throughout this effort. Financial support across the landscape of some ephemeral trees or stand for this project was provided by U.S. Forest Service, conditions [5, 37]. At the landscape scale in central Appa- Northern Research Station. lachian oak-pine forests, prescribed fire has been shown capable of providing that stand condition heterogeneity [38]. 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Journal

International Journal of Forestry ResearchHindawi Publishing Corporation

Published: Jul 2, 2021

References