Geographic Range
Rafinesque's big-eared bats
Corynorhinus rafinesquii
are endemic to North America and are found throughout most of the south-central and
southeastern United States. Their range extends no further north than 40.9 degrees
north in Illinois, Indiana, and Ohio. In the east, its range extends to 37.0 degrees
north is found no further north than southern Virginia. The range extends eastward
to the Atlantic coast and westward to southeastern Texas just east of Dallas, at about
96.6 degrees west. There are two subspecies of the Rafinesque's big-eared bat,
Corynorhinus rafinesquii rafinesquii
and
Corynorhinus rafinesquii macrotis
.
Corynorhinus rafinesquii rafinesquii
is mainly found in the northern and western reaches of the range, whereas
Corynorhinus rafinesquii macrotis
is found in the southeastern states.
Habitat
The habitat requirements of
Corynorhinus rafinesquii
are dependent on the availability of foraging habitat and roost habitat. Foraging
habitat includes mature bottomland hardwoods, young pine
Pinus
stands, oak-hickory (
Quercus
-
Carya
) forests, open field edges, and riparian habitats such as swamps and stream edges.
Foraging habitats must contain a high diversity of woody plants in order to satisfy
the needs of their
lepidopteran
prey.
The subspecies
Corynorhinus rafinesquii rafinesquii
is found in the northern region of the range and commonly forages in mature hardwood
habitats along ridgelines and corridors. The subspecies
Corynorhinus rafinesquii macrotis
inhabits more Coastal Plain foraging habitats, focusing on upland pine stands. This
subspecies will also selectively forage in mature oak and hickory forests. For unknown
reasons, yellow polar (
Liriodendron tulipifera
), American beech (
Fagus grandifolia
), and maple (
Acer
) stands are avoided by both subspecies when foraging.
The roost habitat of the Rafinesque’s big-eared bat can be further broken into three
categories: hibernating roosts, day roosts, and maternity roosts. Roost habitats mainly
consist of caves, dead or decaying trees, and human structures such as barns, houses,
culverts, mine shafts, and bridges. All types of roosts are dependent upon temperature
stability. Most human structures occupied by Rafinesque’s big-eared bats are abandoned
due to a high sensitivity to human disturbance. A roost temperature of 10 to 15 degrees
Celsius is required year-round. Maternity roosts are used from late April to mid-July
and are used for rearing pups. Reproductively active females and their young only
occupy these roosts. Maternity roosts can consist of caves, snags, or human structures.
Day roosts of female Rafinesque’s big-eared bats are used throughout early spring
and early fall (March-April and August-October). For males, day roosts are used consistently
throughout March-October. Day roosts are normally comprised of snags, under tree bark,
in barns, under bridges, in mine shafts, in culverts, and occasionally in caves. Hibernating
roosts, or hibernacula, are used throughout the cold winter months (approximately
November to February). Ideal hibernacula for Rafinesque’s big-eared bat are caves
or mine shafts that maintain a constant temperature of about 11 degrees Celsius. In
the southern portion of the range, hibernacula have included human structures as well
as snags. Roost choice is dependent upon proximity to a water source as well as foraging
habitat. Rafinesque’s big-eared bats are not known to travel much further than 2.5
km from the roost site. Studies have shown that they demonstrate roost fidelity from
year-to-year for all types of roost habitat.
- Habitat Regions
- temperate
- terrestrial
- Other Habitat Features
- suburban
- agricultural
- riparian
- caves
Physical Description
Corynorhinus rafinesquii has relatively flat, dull fur and is brownish-gray in color, sometimes with a reddish hue on the dorsal side. The ventral fur is much lighter, with fur that is dark at the base and light at the tip. This bat is about 10.5 cm in total snout-vent length. The toe hairs extend noticeably beyond the toe. This bat is known for its large ears, which are about 3 cm long in adults. A long, pointed tragus is present in the inner ear that is roughly two-thirds the total ear length. Rafinesque’s big-eared bats have the ability to curl their ears back, which make the ear resemble ram's horns.
There is no difference in appearance between the two subspecies of Rafinesque’s big-eared bat. An additional cusp is present on the upper incisor of Rafinesque's big-eared bat that is not seen in other species in the genus Corynorhinus . The dental formula is: 4/6 2/2 4/6 6/6.
Adults can weigh from 7 to 13 grams and demonstrate sexual dimorphism. Females are normally about one gram larger than adult males. Newborn pups are born naked, without fur and weigh between two to three grams. Neonates will resemble adults in appearance within three weeks of birth.
Large, protruding pararhinal glands are present on the upper side of the snout with extended nostril openings. Rafinesque’s big-eared bats have a wingspan of approximately 25 to 30 cm. Forearm length is between 38 and 45 mm in adults.
Rafinesque’s big-eared bats closely resemble Townsend’s big-eared bats (
Corynorhinus townsendii
) but are distinguishable by several key characteristics. The main difference between
the two is the presence of the additional cusp on the upper incisor of the Rafinesque’s
big-eared bat. There is also a slight difference in fur color between the two species.
Townsend’s big-eared bats tend to have relatively lighter dorsal coat and less variation
in ventral fur color than Rafinesque’s big-eared bats.
- Other Physical Features
- endothermic
- heterothermic
- bilateral symmetry
- Sexual Dimorphism
- female larger
Reproduction
Corynorhinus rafinesquii
mates from late October to late November, before hibernating. Mating occurs during
fall swarms in which males and females gather outside of cave entrances and commence
mating. Female Rafinesque's big-eared bats are polygynandrous and delay fertilization
until after hibernation. Exact length of gestation is unknown. Not much has been published
on the mating systems of
Corynorhinus rafinesquii
. Only hypotheses about the systems have been reported in the literature. For example,
Clark (1990; cited in Lacki and Bayless 2013) suggested that this species has a polygynous
mating system. During the months of fall swarm, she hypothesized that males defend
a territory and encourage multiple non-territorial females to mate with them. Rafinesque’s
big-eared bats are believed to become reproductively active after one year, but little
data are available on reproductive development. The reproductive status of male Rafinesque’s
big-eared bats is distinguishable by the presence of swollen testicles.
- Mating System
- polygynandrous (promiscuous)
Rafinesque’s big-eared bats use internal fertilization. Females form maternity colonies
during late May and early June. During this time females give birth to a single naked
pup, usually weighing 2.3 to 2.6 grams. Females give birth to neonates while roosted
in an upright position, dropping the newborn pups into the membrane between their
hind limbs known as the uropatagium. These bats are thought to be able to distinguish
their young by chemoreception of unique odors produced by newborn pups. Offspring
generally have darker pelage within the first three weeks of life that molts into
adult pelage within eight weeks in southern parts of their range, and 10 to 12 weeks
in the northern parts of their range. Within 4 to 5 weeks after birth, these bats
resemble adults in size. On average, Rafinesque’s big-eared bats become sexually active
around one year of age.
- Key Reproductive Features
- iteroparous
- seasonal breeding
- gonochoric/gonochoristic/dioecious (sexes separate)
- sexual
- viviparous
- sperm-storing
- delayed fertilization
Rafinesque’s big-eared bat young are altricial. These pups are dependent upon maternal
care for three to four weeks during which time they rely on nutrition from their mother’s
milk. Upon the loss of milk teeth, around four weeks of age, these bats gain permanent
dentition and forgo suckling. Young Rafinesque’s big-eared bats begin practicing flight
within three weeks of birth and become competent in flight around five weeks. Oftentimes,
bat captures in late July and early August occur in pairs of mothers and juveniles.
Such observations have led to speculation that there may be a teaching period in which
mothers forage with their young in order to demonstrate proper foraging techniques.
- Parental Investment
- altricial
- female parental care
-
pre-weaning/fledging
-
provisioning
- female
-
protecting
- female
-
provisioning
-
pre-independence
-
provisioning
- female
-
protecting
- female
-
provisioning
Lifespan/Longevity
Corynorhinus rafinesquii
has a lifespan that depends on several factors, including social behaviors, roost
selection, and geographic location. The longest recorded lifespan was of a female
captured in West Virginia that was ten years and one month old (Paradiso and Greenhall,
1967). Little data are available on the lifespan of this bat in the wild.
Most bats can only be tracked via band recoveries, and Lacki (2013) acknowledges that
most banded individuals aren’t recaptured after three years. There are speculations
that banded individuals may avoid recapture by relocating, or becoming savvy to avoid
nets. There are no scientific sources reported on captivity of this bat, although
similar species (e.g.,
Corynorhinus townsendii virginianus
, Virginia big-eared bats) perish at an accelerated rate in captivity. In 2010 the
National Zoo took forty Virginia big-eared into captivity for ensured preservation
of the subspecies, within four months only eleven bats remained. The zoo found that
the most substantial challenge with keeping these bats in captivity was to stimulate
feeding behaviors within a confined area.
Behavior
Seasonal behaviors of Corynorhinus rafinesquii can be broken into three categories, fall swarming and mating, maternity colony formations and pup rearing, and hibernation. Mixed sex roosts are occupied throughout the early spring (March through April), and fall-winter months (August through February). Both sexes occupy day roosts as well as hibernacula together. Rafinesque’s big-eared bats cluster together when roosting and can vary in number of individuals from 2 to 100, while some remain solitary. It is hypothesized that cluster size is inversely related to roost temperature. While roosting, Rafinesque’s big-eared bats hang upside down with the ventral side resting along a substrate. These bats demonstrate grooming behaviors while roosting, which is believed to strengthen social bonds amongst conspecifics. There is no preference in roost choice between sexes. During the summer months (May to August), reproductively-active females gather in maternity colonies and care for their offspring. Roost fidelity is often seen in maternity roosts and these will be used for multiple seasons, upwards of 5 to 10 years, depending on structural integrity of the roost. Maternity roosts can consist of several roost locations including roosts in dead trees, under bridges, and in abandoned human structures. Maternity roost structure is generally comprised of a main home roost with several other satellite roosts, depending on habitat quality and roost availability. Females fly with the young clinging to their ventral side in order to relocate to different roosts. This behavior does not occur during foraging. Males and non-reproductive females continue to use day roosts throughout the summer months. During the fall months (September to October), Rafinesque’s big-eared bats gather in fall swarms and begin to mate. These swarms usually occur outside of cave entrances where mixed sex roosts are located. Rafinesque’s big-eared bats usually do not share roosts with other bat species. However, in the northern parts of the range these bats may occasionally roost with tri-colored bats ( Perimyotis subflavus ) and Virginia big-eared bats ( Corynorhinus townsendii virginianus ). Rafinesque’s big-eared bats hibernate for as long as three to four months in the winter (November to February) to endure cold temperatures. Length of hibernation can vary depending on location and severity of weather from year to year. In the northern portions of their range, Rafinesque’s big-eared bats may undergo a state of torpor during summer months. These bats have the ability to initiate immediate flight from a stationary position if aroused. Although while hibernating it may take several minutes for bats to initiate flight if aroused.
Rafinesque’s big-eared bats usually leave the roost well after dark and return in
the early morning before twilight hours. They are versatile flyers and are able to
travel through cluttered corridors as well as hover in order to capture stationary
prey.
- Key Behaviors
- troglophilic
- flies
- nocturnal
- motile
- hibernation
- daily torpor
- social
- colonial
Home Range
Home range varies depending on location and habitat quality although Menzel et al.
(2001) reported the average home range of 5 male Rafinesque’s big-eared bats in South
Carolina was 93.1 ha (range 61.6 to 225.3 ha). Little literature is available for
the size of the home range in females; it is speculated that it is smaller than males
and more variable with the season.
There are little data available on territory maintenance of these bats and there is
likely overlap between individual territories.
Communication and Perception
Rafinesque's big-eared bats use a low frequency call during echolocation, compared
to many bat species in the genus
Myotis
. All calls are less than ten milliseconds in length and can be described as a frequency-modulated
sweep. The range of the call during the sweep usually begins at about 40 kHz and ends
around 20 kHz. The nature of the low frequency call makes it difficult to record on
acoustic detectors and is believed to aid in the location of stationary prey.
While in flight, this bat continuously has its mouth open in order to produce echolocation
calls. Contrary to popular belief, this bat also uses its eyesight while in flight
in order to avoid obstructions. The elongated nostrils of Rafinesque's big-eared
bats are said to aid the production of ultrasonic sounds, which is thought to help
individuals maintain spatial distribution during foraging. Their large ears also aid
in what is known as passive listening in order to locate stationary prey. Rafinesque's
big-eared bats also have the ability to produce social calls. Although not much information
is available on the diversity of social calls for this species, it is thought that
several types of social calls are used, including alarm calls, mating calls, and territorial
calls. Chemical signals and tactile communication are also suspected to aid in offspring
recognition and roost location. There is no known difference in communication and
perception between subspecies.
- Perception Channels
- visual
- tactile
- acoustic
- ultrasound
- echolocation
- chemical
Food Habits
Rafinesque's big-eared bats are versatile flyers and are able to capture prey by gleaning as well as aerial hawking. The ability to hover and passively listen allows them to be especially successful in the capture of stationary prey species. Rafinesque's big-eared bats forage about one meter off the ground in locations with plenty of vertical and horizontal surface that is necessary for gleaning. The average area of foraging habitat for Rafinesque's big-eared bats is 137.4 hectares and may be occupied by several individuals. Conspecifics are able to maintain a constant spatial distribution during foraging by picking up one another’s echolocation calls and avoiding overlap. Rafinesque's big-eared bats use feeding perches, where they consume prey that they have caught. A late flyer, they usually leave the roost to forage several hours after dark. Males may demonstrate biphasic foraging patterns using two different foraging trips a night. The first being from about an hour after dark until midnight and the second beginning around four in the morning until just before sunrise.
The areas used for foraging can differ in individuals based on time of year, sex, and reproductive condition. Rafinesque’s big-eared bats are forest habitat generalists and target areas along forest edges, along cliff walls, and in riparian corridors.
Rafinesque's big-eared bats are Lepidopteran specialists. Studies have shown that
more than 80% of the diet consists of various
lepidopteran
species including six families of moths and 22 species. Wingspan of lepidopteran
prey is a limiting factor. The average wingspan of lepidopteran prey is between 47
and 70 millimeters. The family
Noctuidae
makes up the bulk of the diet at about 39 percent. The five other main families include
Arctuidae
,
Geometridae
,
Megalopygidae
,
Notodontidae
, and S<
phingidae
>. Studies have shown that Rafinesque's big-eared bats also forage on beetles (
Coleoptera
), cicadas (
Hemiptera
), flies (
Diptera
), true bugs (
Hemiptera
), bees and ants (
Hymenoptera
), and caddis flies (
Trichoptera
). There is no evidence of difference in food habits between subspecies.
- Primary Diet
- carnivore
- Animal Foods
- insects
Predation
Due to the nocturnal nature and versatile flying abilities of Rafinesque’s big-eared
bats, there are few known predators that are successful at capturing these bats while
in flight, most predation occurs while bats are roosted. Bats that choose less cryptic
roost locations such as abandoned buildings tend to be more susceptible to predation.
There are few reported cases of observed predation of these bats however black rat
snakes (
Pantherophis alleghaniensis
), raccoons (
Procyon lotor
), Virginia opossums (
Didelphis virginiana
), and house cats (
Felis catus
) are known predators of this species. In some cases young Rafinesque’s big-eared
bats may become dislodged from their roosts and fall to the ground leaving them vulnerable
to predators, including snakes and terrestrial predators.
- Anti-predator Adaptations
- cryptic
Ecosystem Roles
This bat is a lepidopteran specialist and significantly controls insect populations
throughout its range. Rafinesque’s big-eared bats host both ectoparasites and endoparasites.
A study by McAllister et al. (2005) reported a variety of gastrointestinal helminths
present in 10% of the observed bats in Arkansas. The three main parasites recorded
in this study include one parasite in the class Cestoda (
Vampirolepis
) and two parasites from the phylum Nematoda (
Physaloptera
and
Capillaria palmata
). These specific parasites are common to occur in the stomach, small intestine, and
feces of the Rafinesque’s big-eared bat. Rafinesque’s big-eared bats host a variety
of ectoparasites that may be present on exposed patagium as well as in the pelage.
Little data are available on the multitude of external parasites of this bat although
several species of mites, fleas, flies, and ticks are believed in infect this bat.
- cestodes ( Vampirolepis )
- nematodes ( Physaloptera )
- nematodes ( Capillaria palmata )
Economic Importance for Humans: Positive
Like most North American bat species,
Corynorhinus rafinesquii
is an insectivorous bat that potentially plays a role in pest control throughout
its range. The presence of this bat directly contributes to the population control
of several insect species groups, including mainly lepidopteran moth species. Lepidopteran
moths tend to lay their eggs on woody shrubs which provide nourishment for the young
while at the same time harm the woody plant species. Valuable tree species including
those from the genus
Pinus
,
Quercus
, and
Carya
can be affected by these moths. A study by Boyles et al. (2011) predicted that the
disappearance of bats in North America could lead to an estimated $3.7 billion/year
in agricultural losses.
- Positive Impacts
- controls pest population
Economic Importance for Humans: Negative
Corynorhinus rafinesquii
is a known carrier of the rabies virus and a fungal disease known as histoplasmosis
that can be contracted by humans. No definitive figures on confirmed cases of the
rabies virus being present in this bat are available. As a species that roosts in
abandoned human structures, fecal accumulation containing pathogens may be hazardous
to humans. However, the chance of such transference is very low. Klug et al. (2011)
reported that in a study on the prevalence of rabies in bats, five Rafinesque’s big-eared
bats were tested for the rabies virus and zero bats tested positive for the virus.
- Negative Impacts
-
injures humans
- carries human disease
Conservation Status
Currently, Corynorhinus rafinesquii is listed as a species of least concern on the IUCN red list, although previous (1996) listings report it as vulnerable. afinesque’s big-eared bats are sensitive to habitat alteration and human disturbance. With human expansion and development increasing at an alarming rate, the most detrimental impact on this species is habitat loss. An increase in logging practices that destroy roost habitats in the range of these bats poses a serious threat. This bat relies on hardwood habitats in order to maintain summer roosting habitat and foraging habitat. The main prey item of the Rafinesque’s big-eared bat is moths, which may rely on mature woody plants for larval development. These woody plants such as oak ( Quercus ), poplar ( Populus ), maple ( Acer ), and pine ( Pinus ) are commonly targeted for timber harvests.
Another concern for this species is the destruction of roost locations. In some cases, these bats roost in various human structures, such as abandoned buildings, mine shafts, and bridges. Human activity may destroy active or potential roosts. Human disturbance during hibernation periods can cause harm to these bats. Conservation strategies prohibiting human entrance into hibernacula are suggested to avoid disturbance and arousal of this species during hibernation.
Since the discovery of white-nose syndrome (WNS) in New York in 2006, populations
of some bats that use cave hibernacula in eastern North America have been decimated.
Rafinesque’s big-eared bats, as well as other bat species in the genus
Corynorhinus
, have not demonstrated susceptibility to the fungus that causes WNS. Bernard et al.
(2015) reported the first confirmed case of
Pseudogymnoascus destructans
, the fungus that causes WNS, present on the skin of two individuals outside of a
cave in Tennessee. Johnson et al. (2012) suggested that species-specific winter activity
patterns such as frequent arousal from torpor along with other ecological and physiological
adaptations provide defense against the harmful effects of WNS.
Additional Links
Contributors
Keifer Titus (author), Radford University - Fall 2015, Cari Mcgregor (editor), Radford University, Zeb Pike (editor), Radford University, Karen Powers (editor), Radford University, April Tingle (editor), Radford University, Jacob Vaught (editor), Radford University, Tanya Dewey (editor), University of Michigan-Ann Arbor.
- Nearctic
-
living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- temperate
-
that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).
- terrestrial
-
Living on the ground.
- forest
-
forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.
- mountains
-
This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.
- marsh
-
marshes are wetland areas often dominated by grasses and reeds.
- swamp
-
a wetland area that may be permanently or intermittently covered in water, often dominated by woody vegetation.
- bog
-
a wetland area rich in accumulated plant material and with acidic soils surrounding a body of open water. Bogs have a flora dominated by sedges, heaths, and sphagnum.
- suburban
-
living in residential areas on the outskirts of large cities or towns.
- agricultural
-
living in landscapes dominated by human agriculture.
- riparian
-
Referring to something living or located adjacent to a waterbody (usually, but not always, a river or stream).
- endothermic
-
animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.
- heterothermic
-
having a body temperature that fluctuates with that of the immediate environment; having no mechanism or a poorly developed mechanism for regulating internal body temperature.
- bilateral symmetry
-
having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.
- polygynandrous
-
the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.
- iteroparous
-
offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).
- seasonal breeding
-
breeding is confined to a particular season
- sexual
-
reproduction that includes combining the genetic contribution of two individuals, a male and a female
- viviparous
-
reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.
- sperm-storing
-
mature spermatozoa are stored by females following copulation. Male sperm storage also occurs, as sperm are retained in the male epididymes (in mammals) for a period that can, in some cases, extend over several weeks or more, but here we use the term to refer only to sperm storage by females.
- delayed fertilization
-
a substantial delay (longer than the minimum time required for sperm to travel to the egg) takes place between copulation and fertilization, used to describe female sperm storage.
- altricial
-
young are born in a relatively underdeveloped state; they are unable to feed or care for themselves or locomote independently for a period of time after birth/hatching. In birds, naked and helpless after hatching.
- female parental care
-
parental care is carried out by females
- nocturnal
-
active during the night
- motile
-
having the capacity to move from one place to another.
- hibernation
-
the state that some animals enter during winter in which normal physiological processes are significantly reduced, thus lowering the animal's energy requirements. The act or condition of passing winter in a torpid or resting state, typically involving the abandonment of homoiothermy in mammals.
- social
-
associates with others of its species; forms social groups.
- colonial
-
used loosely to describe any group of organisms living together or in close proximity to each other - for example nesting shorebirds that live in large colonies. More specifically refers to a group of organisms in which members act as specialized subunits (a continuous, modular society) - as in clonal organisms.
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- chemical
-
uses smells or other chemicals to communicate
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- ultrasound
-
uses sound above the range of human hearing for either navigation or communication or both
- echolocation
-
The process by which an animal locates itself with respect to other animals and objects by emitting sound waves and sensing the pattern of the reflected sound waves.
- chemical
-
uses smells or other chemicals to communicate
- cryptic
-
having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.
- carnivore
-
an animal that mainly eats meat
- insectivore
-
An animal that eats mainly insects or spiders.
References
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Bernard, R., J. Foster, E. Willcox, K. Parise, G. McCracken. 2015. Molecular detection of the causative agent of white-nose syndrome on Rafinesque's big-eared bats (Corynorhinus rafinesquii) and two speceis of migratory bats in the southeastern USA. Journal of Wildlife Diseases , 51/2: 519-522.
Boyles, J., P. Cryan, G. McCracken, T. Kunz. 2011. Economic importance of bats in agriculture. Science , 332/6025: 41-42.
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Hurst, T., M. Lacki. 1997. Food habits of Rafinesque's big-eared bat in southeastern Kentucky. Journal of Mammalogy , 78/2: 525-528.
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Johnson, J., J. Kropczynski, M. Lacki, G. Langlois. 2012. Social networks of Rafinesque's big-eared bat (Corynorhinus rafinesquii) in bottomland hardwood forests. Journal of Mammalogy , 93/6: 1545-1558.
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