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2009 SOUTHEASTERN NATURALIST 8(2):355–362
Myotis septentrionalis Trouessart (Northern Long-eared
Bat) Records from the Coastal Plain of North Carolina
Adam D. Morris1, Maarten J. Vonhof2, Darren A. Miller3,
and Matina C. Kalcounis-Rueppell1,*
Abstract - Myotis septentrionalis (Northern Long-eared Bat) is a small, insectivorous
bat found in the eastern United States and Canada. Along the east coast, its range
is thought to extend as far south as the Great Dismal Swamp in coastal Virginia. We
captured six M. septentrionalis in the northern coastal plain region of North Carolina.
Field identification was based on characters of ear and tragus length and confirmed
with mitochondrial cytochrome c oxidase subunit I sequences. These captures signify
the presence of a resident population of M. septentrionalis in the northern coastal
plain of North Carolina. Future work is needed to document range limits and hibernation
behavior of this species in the piedmont and coastal plain of North Carolina.
Myotis septentrionalis Trouessart (Northern long-eared Bat; Fig 1.) is a
small insectivorous bat found in the eastern United States and Canada. Myotis
septentrionalis was formerly thought to be a subspecies of M. keenii
(Merriam) (Keen’s Myotis; Miller and Allen 1928), but is now recognized
as a distinct species (van Zyll de Jong 1979). Myotis septentrionalis is
distinguished from other Myotis species by its large ear, that, when laid
forward, extends beyond the muzzle and its long, pointed tragus (Caceres
and Barclay 2000).
Myotis septentrionalis hibernates in caves and abandoned mines during
the winter, where it is commonly observed swarming (Caceres and Barclay
2000). Copulation occurs in hibernacula prior to winter, and females store
sperm and fertilize a single egg in the spring. During the summer months,
M. septentrionalis uses trees as day-roosts (Caceres and Barclay 2000).
Myotis septentrionalis is a small and maneuverable bat that hunts within
cluttered forest stands (Caroll et al. 2002, Owen et al. 2003) in both upland
(Harvey et al. 1999, Lacki and Schwierjohann 2001, Sasse and Pekins 1996)
and fl oodplain forests (Carroll et al. 2002, Foster and Kurta 1999). Myotis
septentrionalis is known to hunt fl ying insects and also glean insects from
substrates (Faure et al. 1993).
Myotis septentrionalis occupies much of the eastern United States and
Canada, but is most common within Ontario, Quebec, and the New England
states (Caceres and Barclay 2000). However, its range extends south into
1Biology Department, University of North Carolina at Greensboro, Greensboro, NC
27403. 2Department of Biological Sciences, Western Michigan University, Kalamazoo,
MI 49008. 3Southern Timberlands Research and Development, Weyerhaeuser
Company, Columbus, MS 39704. *Corresponding author - firstname.lastname@example.org.
356 Southeastern Naturalist Vol.8, No. 2
Alabama and Georgia, and west into Alberta, British Columbia, Montana,
and Wyoming (Caceres and Barclay 2000), and the species can be locally
common in these regions. Along the east coast, the range is thought to extend
as far south as the Great Dismal Swamp in coastal Virginia (specimens:
AMNH 93177 and USNM 23277, see Appendix 1 for full names; Webster
et al. 1985). Previous distribution maps suggesting a presence in the North
Carolina coastal plain (Caceres and Barclay 2000) are drawn broadly and
are based on few actual records. Although the Great Dismal Swamp extends
into North Carolina, recent surveys have found no evidence of M. septentrionalis
in the coastal plain of North Carolina (Clark 1999, Lambiase et al.
2002, McDonnell 2001). Records of M. septentrionalis in North Carolina
are mainly from the western portion of the state, with the exception of one
isolated record in the piedmont (NCSM 45; Fig. 2) and one reportedly in the
southern coastal plain (David Webster, University of North Carolina, Wilmington,
NC, pers. comm.; Fig. 2).
During the summer of 2007, we captured bats using mist-nets at the Tidewater
Research Station, located in the coastal plain 5 miles east of Plymouth
in Washington County, NC. We captured six M. septentrionalis in mist-nets
set along a closed-canopy, overgrown road corridor directly adjacent to a
natural forested wetland area surrounded by intensively managed Pinus taeda
L. (Loblolly Pine). Field identification to species included measurements
Figure 1. Individual juvenile female M. septentrionalis (ID ADM35 from Table 1)
captured in the northern coastal plain of North Carolina during summer 2007.
2009 A.D. Morris, M.J. Vonhof, D.A. Miller, and M.C. Kalcounis-Rueppell 357
and characters of ear and tragus length (Table 1; Fig 1). We also collected
tissue samples from wing membranes, and all bats were released at site of
capture. We did not mark captured bats in any way. However, because we
collected a wing biopsy, we could tell if individuals were recaptures based
on either the presence of the biopsy site or scaring over the biopsy site.
Capture and handling protocols were approved by the UNCG Institutional
Figure 2. Map showing the range of M. septentrionalis in North Carolina, South
Carolina, Virginia, and Maryland. Shaded counties contain documented records of M.
septentrionalis; the filled black circle represents our capture site. Map was generated
using a map layer compiled by Bat Conservation International (available at http://
nationalatlas.gov/mld/bat000m.html) and a compilation of local records (D. Webster,
Table 1. Measurements of the six individuals of M. septentrionalis captured in the northern
coastal plain of North Carolina during summer, 2007. Measurements of length were taken with
standard metric calipers and measurements of mass taken with a Pesola® spring scale.
Forearm Ear Weight
ID Date Time Sex Age (mm) (mm) (g)
ADM31 6/11/2007 10:15 PM Male Adult 33 n/a 4.95
ADM34 6/18/2007 9:38 PM Female Juvenile 34 16 4.95
ADM35 6/18/2007 11:40 PM Female Juvenile 34 16 5.45
ADM36 6/25/2007 9:50 PM Female Juvenile 36 n/a 5.95
ADM41 6/25/2007 2:15 AM Male Juvenile 36 15 5.50
ADM50 7/2/2007 10:05 PM Female Juvenile 35 n/a 5.45
358 Southeastern Naturalist Vol.8, No. 2
Animal Care and Use Committee (# 06-11) and complied with recommendations
of the American Society of Mammalogists.
To confirm species identification, we compared mitochondrial cytochrome
c oxidase subunit I (COI) sequences from these six bats to those of
M. septentrionalis collected elsewhere and to other eastern North American
bats in the genus Myotis that are also found in North Carolina (Fig. 3, Appendix
2). We obtained partial COI sequences (bidirectionally sequenced)
using primers and cycling conditions outlined in Hebert et al. (2003), and
cleaned and aligned sequences using CodonCode Aligner 2.04 (CodonCode
Corporation, Dedham, MA), resulting in a final fragment length of 636 bp.
We estimated sequence divergences by using the Kimura-2-Parameter distance
model and graphically displayed these in a neighbor-joining tree using
PAUP (v. 4.0b10; Swofford 2002).
Figure 3. Neighbor-joining phylogram of COI sequences based on Kimura 2-parameter
distances, showing the specimens from Washington County, NC (two unique
haplotypes among the six specimens, labeled ADM31 and ADM50) grouping with
other M. septentrionalis samples from other parts of the range. Details of specimens
included can be found in Appendix 2.
2009 A.D. Morris, M.J. Vonhof, D.A. Miller, and M.C. Kalcounis-Rueppell 359
Results and Discussion
Five of the six bats captured in North Carolina shared identical
haplotypes (ADM31 through ADM41; see Table 1, Figure 3). Four M.
septentrionalis specimens (CM82047, CS04, JJ43, UAM68932) from various
locations shared the same haplotype (Fig. 3). The two M. austroriparius
specimens (FBF11 and AM168) shared the same haplotype (Fig. 3). Mean
sequence divergence between the two unique haplotypes observed in North
Carolina and other M. septentrionalis specimens (4 unique haplotypes) was
0.4%, while sequence divergence with other eastern North American Myotis
averaged 9.7% (ranging from 8.5% with M. lucifugus to 13.8% with M. austroriparius).
The two haplotypes from North Carolina clearly grouped with
other M. septentrionalis in the neighbor-joining analysis (Fig. 3), confirming
identification of the six bats.
The six bats we captured represent both adult and juveniles (young of the
year in 2007), suggesting reproduction of these bats in this area. These six
bats were captured mid-breeding season (11 June through 2 July) on 4 different
nights at the same mist-net site, suggesting that these were not migrating
individuals. Thus, based on seasonal timing of adult and juveniles captured
over multiple nights, we suggest that these M. septentrionalis signify a
resident population of this species in the northern coastal plain of North
Carolina as opposed to stray captures.
This population is 96 km further south of the Great Dismal Swamp localities
in Virginia. The area where these bats were captured refl ects habitat
preferences of this species in other parts of its range (Carroll et al. 2002,
Harvey et al. 1999, Owen et al. 2003). That is, these bats were captured in a
mist-net strung across an overgrown roadway adjacent to mature forest stands.
However, our captures were not in close proximity to cave hibernacula. Myotis
septentrionalis are thought remain close to hibernacula during summer foraging
(Caceres and Barclay 2000). Since no caves exist in the coastal plain of
North Carolina, our captures imply that these bats either 1) travel much further
to hibernacula than previously thought; 2) utilize alternative hibernacula, such
as tree cavities; or 3) do not hibernate during the mild winters in this area. In
the coastal plain, M. septentrionalis may hibernate in tree cavities or buildings,
like Corynorhinus rafinesquii (Lesson) (Rafinesque’s Big-eared Bat)
(Trousdale and Beckett 2005, Trousdale et al. 2008).
Future work is needed to document range limits of this species and to
fill in gaps between the location in our study and known locations of the
species to the north and west. Future studies should also examine hibernation
behaviors of M. septentrionalis in the piedmont and coastal plain of
Funding was provided by Weyerhaeuser Company, the University of North Carolina
at Greensboro, and Western Michigan University. J. Hart-Smith and D. Allgood
assisted with fieldwork. Discussions with D. Webster, M. Clark, and L. Gatens were
360 Southeastern Naturalist Vol.8, No. 2
appreciated. We thank the curators and institutions for their loans of specimens and
tissues: L.K. Ammerman with the Angelo State Natural History Collection, S.B.
McLaren with the Carnegie Museum of Natural History, and L. Olson with the University
of Alaska Museum. We are also indebted to the researchers who provided
wing-membrane samples, including R. Benedict, E. Britzke, M. Clark, R. Currie, E.
Gates, J. Johnson, A. Miles, and C. Stihler.
Caceres, M.C., and R.M.R. Barclay. 2000. Myotis septentrionalis. Mammalian Species
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Effects on perceived fauna. Southeastern Naturalist 1:193–198.
Clark, M.K. 1999. Results of a bat survey in the lower Roanoke River Basin. Report
prepared for the North Carolina Natural Heritage Trust Fund, North Carolina
Natural Heritage Program, Raleigh, NC. 36 pp.
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Long-eared Bat, Myotis septentrionalis, are relatively inaudible to moths.
Journal of Experimental Biology 178:173–189.
Foster, R.W., and A. Kurta. 1999. Roosting ecology of the Northern Bat (Myotis septentrionalis)
and comparisons with the endangered Indiana Bat (Myotis sodalis).
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Bats in mixed mesophytic forest. Journal of Wildlife Management 65:482–488.
Lambiase, S.J., M.K.Clark, and L.J. Gatens. 2002. Bat (Chiroptera) Survey of North
Carolina State Parks 1999–2001. North Carolina Division of Parks and Recreation
and North Carolina State Museum of Natural Sciences, Raleigh, NC.
McDonnell, J.M. 2001. Use of bridges as day roosts by bats in the North Carolina
coastal plain. M.Sc. Thesis. North Carolina State University, Raleigh, NC. 74 pp.
Miller, G.S., Jr., and G.M. Allen. 1928. The American bats of the genera Myotis and
Pizonyx. Bulletin of the United States National Museum 144:1–218.
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(Myotis septentrionalis). American Midland Naturalist 150:352–359.
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362 Southeastern Naturalist Vol.8, No. 2
Appendix 1. Full names of museum acronyms for specimens listed in text and not
used in the genetic analysis.
AMNH = American Museum of Natural History.
NCSM = North Carolina State Museum of Natural Sciences.
USNM = National Museum of Natural History.
Appendix 2. Locations of specimens used in the genetic analyses. Voucher specimens
used in this study were housed in the Carnegie Museum of Natural History,
Pittsburgh, Pennsylvania (CM), the Angelo State Natural History Collection, San
Angelo, TX (ASK), and the University of Alaska Museum, Fairbanks, AK (UAM).
Additional wing membrane samples were used, and housed at Western Michigan
University (all other abbreviations).
Myotis austroriparius – UNITED STATES. South Carolina: Francis Beidler Forest,
Dorchester County (FBF11). Georgia: Climax Cave, Decatur County (AM168).
Myotis leibii – UNITED STATES. West Virginia: North Fork Mountain, Pendleton
Myotis lucifugus – UNITED STATES. Montana: Flathead National Forest, 0.5 mi
NW of Swan Lake, Lake County (ASK4402). West Virginia: Babcock State Park, 0.2
miles S, 0.3 miles W of Clifftop, Fayette County (CM102862). Nebraska: Guadalcanal
Prairie, 5.7 miles S and 3.5 miles W of Harrison, Sioux County (RB5828).
Myotis septentrionalis – UNITED STATES. Kentucky: Bangor, Rowan County
(EB95). Maryland: Owens Creek campground, Catoctin Mountain Park, Frederick
County (JJ43). Nebraska: Walnut Creek, 3.9 mile N and 4.4 miles W of Newcastle,
Dixon County (RB5610). West Virginia: Monongahela National Forest, 3 miles N,
4.5 miles W of Harmon, Randolph County (CM82047). North Fork Mountain, Pendleton
County (CS04). Camp Creek State Forest and Park, Mercer County (CS11).
CANADA. British Columbia: Smith River, N of Alaska Highway (UAM68932)
Myotis sodalis – UNITED STATES. Tennessee: White Oak Blowhole Cave, Blount
County (MYSO304). Virginia: Cumberland Gap Saltpeter Cave, Lee County