Myotis lucifuguslittle brown bat

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Geographic Range

Little brown bats, Myotis lucifugus, are abundant in southern Alaska, Canada, across the United States from the Pacific to Atlantic coasts, and the higher elevation forested regions of Mexico. Although little brown bats are not found in northern Canada, individuals have been observed in Iceland and Kamchatka. Those outlying records are presumed to have been the result of accidental ship transportation by humans. Little brown bats are also absent from much of Florida, the southern Great Plains regions of the U.S., southern California, and parts of coast Virginia and the Carolinas. (Barbour and Davis, 1969; Fenton and Barclay, 1980; Nowak, 1994)

Habitat

Myotis lucifugus occupies three types of roosts: day, night, and hibernation roosts. Locations of roosts are chosen based upon the presence of stable ambient temperatures. Day and night roosts are used by active bats and include, but are not limited to, buildings, trees, under rocks, and in piles of wood. Day roosts have very little or no light, provide good shelter, and typically have southwestern exposures to provide heat for arousal from daily torpor.

Night roosts are selected for their confined spaces where large concentrations of bats can cluster together to increase the temperature in the roost. These roosts are primarily occupied when temperatures are below 15°C. Night roosts are usually away from day roosts; this may diminish the accumulation of feces at day roosts and avoid signaling predators. Day and night roosts are inhabited during spring, summer, and fall months, whereas during the winter, hibernacula sites are used.

Nursery roosts are similar to day roosts but are warmer than ambient temperature. They are usually occupied only by females and their offspring. Females use the same nursery colony every year.

Hibernaculum sites may be shared with Myotis yumanensis. These sites usually include abandoned mines or caves where the temperature is continuously above freezing and humidity is high. Northern populations of bats enter hibernation in early September and end in mid-May; southern populations enter in November and end their hibernation in mid-March. Myotis lucifugus does not make tremendously long migrations during the change of seasons. (Wilson and Ruff, 1999; Fenton and Barclay, 1980; Koopman and Gudmundsson, 1966; Nowak, 1994; Wilson and Ruff, 1999)

Myotis lucifugus inhabits forested lands near water, but some subspecies can be found in dry climates where water is not readily available. In those habitats, drinking water is provided by moisture on cave walls or condensation on the fur. Little brown bats live over a wide latitudinal and elevational range. (Barbour and Davis, 1969; Fenton and Barclay, 1980; Tuttle, 1991; Wilson and Ruff, 1999)

Physical Description

The fur of M. lucifugus is glossy, and varies in color from dark brown, golden brown, reddish, to olive brown. Albino individuals have also been observed. The ventral side has lighter pelage. The wing and interfemoral membranes are nearly hairless and dark brown or black. The tragus is blunt and of medium height. Their ears usually do not extend past the nose when laid forward. Myotis lucifugus has small ears and large hind feet. The fore and hind limbs have five metapodials. The hind foot has hairs that extend past the toes.

The skull has some distinguishing characteristics. Myotis lucifugus lacks a saggital crest, has a shortened rostrum, 38 teeth, and a upslope profile of the forehead. In addition, the braincase is flattened and subcircular when observed dorsally.

Myotis lucifugus does not possess a keel on the calcar and has a short tibia relative to the length of the hind foot (~55% of the tibial length). Myotis lucifugus lacks choroidal papillae and folded retinas, and therefore does not exhibit eye shine. (Nowak, 1994; Nowak, 1994; Fenton and Barclay, 1980; Kurta, 1995; Nowak, 1994)

Myotis lucifugus weighs between 5 and 14 g. The length varies between 60 and 102 mm, and the wingspan between 222 and 269 mm. The forearm, including claw, measures 33 to 41 mm, and tail length measures 28 to 65 mm. The hind foot is between 8 and 10 mm in length, ears are 11 to 15.5 mm, and the tragus is 7 to 9 mm. Little brown bats fly at speeds as high as 35 km/hour and average 20 km/hour. Females are larger than males, especially during the winter. (Fenton and Barclay, 1980; Nowak, 1994; Wilson and Ruff, 1999)

  • Sexual Dimorphism
  • female larger
  • Range mass
    5 to 14 g
    0.18 to 0.49 oz
  • Range length
    60 to 102 mm
    2.36 to 4.02 in
  • Average length
    87 mm
    3.43 in
  • Range wingspan
    222 to 269 mm
    8.74 to 10.59 in
  • Average basal metabolic rate
    0.051 W
    AnAge

Reproduction

Mating occurs between adult females and adult males; subadult males are not sexually mature until after their first year. Mating occurs in two phases: active and passive. During the active phase, both partners are awake and alert. In the passive phase, active males mate with torpid individuals of both sexes; passive phase mating is approximately 35% homosexual. Mating is random and promiscuous. Females in active phase usually mate with more than one male. In both active and passive phase matings, males mate with multiple females. (Fenton and Barclay, 1980; Wai-Ping and Fenton, 1988)

Swarming at the hibernacula occurs during late summer and fall; activity decreases with lower temperatures. Swarming serves a prenuptial function, along with showing the young suitable hibernation roosts. During late July, bats arriving at the hibernacula are adult males and nonparous females; females and subadults appear in early August. Swarming M. lucifugus may travel large distances, causing mixing of populations from different areas. During the swarming period, little brown bats are receptive to calls of conspecifics. (Fenton and Barclay, 1980; Schowalter, 1980)

Myotis lucifugus has enlarged pararhinal glands during the mating season. Mating occurs when a male mounts a female from the rear. The male may bite the female on her back. Upon female struggle, the male may emit a copulation call to ease the female. Males inseminate females that are are active as well as those that are torpid. Little brown bats delay ovulation and store sperm for about seven months between copulations in the fall and fertilization in the spring. Pups are born and reared in June and July after a 50 to 60 day gestation period. The greatest energy demand on females occurs during lactation and toward the end of pregnancy. Females lose the ability to thermoregulate well when approaching parturition. (Fenton and Barclay, 1980; Wai-Ping and Fenton, 1988)

Normally, bats hang head down; females giving birth reverse their position, so their head is up. Young are born into the interfemoral membrane; only one young is born per year. The pups’ eyes and ears open within hours of birth, and deciduous teeth are fully erupted. Pups must cling to the female’s nipple using their deciduous incisors, large thumbs, and hind feet. The young start hearing at day 2 and develop auditory sensitivity similar to that of an adult by day 13. On approximately day 9.5, pups are able to thermoregulate and in three weeks they are able to fly.

Independence from the mother comes when the pups start to fly and become self-supporting at about 4 weeks of age. Adult weight is attained at about 4 weeks of age as well. Spermatogenesis starts in May and ends in August. (Fenton and Barclay, 1980)

  • Breeding interval
    Both sexes mate more than once per year and produce one young per year.
  • Breeding season
    Mating begins in mid-August during the active phase and continues in passive phase throughout the winter.
  • Range number of offspring
    1 to 2
  • Average number of offspring
    1
  • Average number of offspring
    1
    AnAge
  • Range gestation period
    50 to 60 days
  • Range weaning age
    21 to 28 days
  • Average weaning age
    26 days
  • Average time to independence
    4 weeks
  • Average age at sexual or reproductive maturity (female)
    210 days
  • Average age at sexual or reproductive maturity (female)
    Sex: female
    210 days
    AnAge
  • Average age at sexual or reproductive maturity (male)
    210 days
  • Average age at sexual or reproductive maturity (male)
    Sex: male
    210 days
    AnAge

Mothers nurse their own young and distinguish from other pups by odor and calls. For 18 to 21 days, pups ingest only milk from their mother. Weaning takes place at about three weeks; at this time, the permanent teeth fully erupt and pups start to feed on insects along with the mother's milk. After weaning, the pups have a drop in body weight as they learn to catch insects. It is not clear if mothers bring insects to their young or help to teach them to hunt. However, many female/young pairs are captured together, suggesting that there is some period of supervised learning before independence. Males play no role in parental care. (Fenton and Barclay, 1980)

  • Parental Investment
  • altricial
  • pre-fertilization
    • provisioning
    • protecting
      • female
  • pre-hatching/birth
    • provisioning
      • female
    • protecting
      • female
  • pre-weaning/fledging
    • provisioning
      • female
    • protecting
      • female
  • pre-independence
    • provisioning
      • female
    • protecting
      • female

Lifespan/Longevity

The lifespan of M. lucifugus is extended by their ability to find food and inhabit a variety of roosts. These characteristics allow expansion of their habitat to new ranges, but also contribute to their survival. Myotis lucifugus live approximately 6 to 7 years and often live well beyond 10 years. A 31 year-old male was discovered in southeastern Ontario. Evidence indicates that males tend to live longer than females. Mortality rate is the greatest during the first winter when new pups have considerably less weight than their adult counterparts at the start of hibernation. (Fenton and Barclay, 1980; Nowak, 1994)

Behavior

Little brown bats are primarily nocturnal and emerge from their roosts at dusk. Primary activity occurs about two or three hours after dusk and secondary activity may occur before dawn; most individuals return to the roost by four or five o’clock in the morning. These bats usually enter daily torpor. During the winter, hibernation time depends on altitude and location of the roosts. It usually starts between September and November and ends in March to May. The young remain active longer in the fall to build fat deposits to last the winter. Myotis lucifugus does not migrate long distances for hibernation roosts. Individuals travel only up to 100 miles. This species does not show territoriality at roosts, and large colonies of as many as 300,000 bats have been reported in a single roost. (Barbour and Davis, 1969; Cockrum, 1956; Nowak, 1994)

During hibernation, little brown bats undergo repeated periods of torpor lasting 12 to 19 days, but may remain torpid for as long as 83 days. Signals for the end of hibernation include weather conditions of the area and arousal of neighboring bats. (Fenton and Barclay, 1980; Tuttle, 1991)

Little brown bats vary their body temperatures greatly. These bats can be cooled to 6.5 degrees Celsius and heated to temperatures of 54 degrees Celsius without harm. Myotis lucifugus deposits 13 distinct type of brown fat, allowing individuals to efficiently and rapidly produce heat during arousal from hibernation torpor.

Myotis lucifugus occultus can increase urine concentration in order to better withstand water stress in low humidity environments with limited water supply. Only this subspecies is known to do this, however; most subspecies of M. lucifugus have a poor ability to regulate urine concentration. In general, M. lucifugus usually lives within close proximity to water. (Fenton and Barclay, 1980; Tuttle, 1991)

Myotis lucifugus invests a large amount of time each day grooming. Individuals use their claws to groom the fur, and tongue and teeth to clean their wing membranes. (Fenton and Barclay, 1980)

Home Range

Little brown bats travel several kilometers between day roosts and feeding sites. (Nowak, 1994)

Communication and Perception

Myotis lucifugus produces frequency modulated (FM) calls at 45kHz, their fundamental frequency. These calls last 1 to 5 milliseconds and sweep from 80 to 40 kHz. Cruising bats typically produce 20 calls per second to detect prey and objects. (Fenton and Barclay, 1980; Fenton and Bell, 1979)

Myotis lucifugus alert other bats with non-echolocation calls if they are flying on a collision course during feeding. They emit this call by reducing the frequency of the terminal portion of a sweep call to 25 kHz. Additionally, they may use echolocation calls, visual cues, such as landmarks, and possibly chemical cues to locate roosts; they can find their roosts from 180 miles away. Mother and young communicate through a few, complex vocalizations. There is no information about alarm or distress calls. (Fenton and Barclay, 1980; Kurta, 1995)

Food Habits

Myotis lucifugus is an efficient insect predator, especially when insects are in patches and at close range (approximately less than one meter). Little brown bats, along with many other insectivorous bats, are opportunistic feeders and catch prey by aerial hawking and gleaning tactics. Myotis lucifugus flies faster near the end of the attack, when approaching prey. During gleaning, these bats hover approximately 30 cm from the prey. Myotis lucifugus typically feeds on swarms of insects, saving time and energy to search for food. There is no evidence of territorial protection of feeding areas, but individuals return to areas where they have had prior feeding success. Myotis lucifugus has different selectivity based upon the arrangements of insects. In large swarms of mating insects, these bats concentrate one or two species to feed on, but when insects are dispersed, little brown bats are less selective and feed on multiple species. Food demand of lactating females increases and pregnant or lactating females usually select larger insects than males or nonpregnant females. Normally, these bats feed on insects whose length ranges from 3 to 10 mm. These bats typically eat half of their body weight per night (when active) and lactating females eat approximately 110 percent of their body weight per night. Myotis lucifugus chews and processes food relatively quickly. Mastication rate is seven jaw cycles per second, and food takes only 35 to 54 minutes to pass items through the digestive system. (Anthony and Kunz, 1977; Belwood and Fenton, 1976; Fenton and Barclay, 1980; Ratcliffe and Dawson, 2003; Wilson and Ruff, 1999)

Myotis lucifugus uses FM echolocation, downward sweeping pulses of 80 to 40 kHZ that last from 1 to 5 msec. These wavelengths give the greatest quality of detection for 3 to 8 mm size prey, and M. lucifugus consumes prey averaging 3 to 10 mm in size. The same FM call is used for location of both flying and stationary insects. The approach phase of their call has second and third harmonics, but during the feeding buzz, the frequency is focused at 47kHz. The rate of call production while chasing prey is 200 calls per second. Myotis lucifugus emits a high-pulse repetitive call when nearing a landing site. (Fenton and Barclay, 1980; Anthony and Kunz, 1977; Barbour and Davis, 1969; Fenton and Barclay, 1980; Fenton and Bell, 1979; Ratcliffe and Dawson, 2003; Wilson and Ruff, 1999)

Myotis lucifugus catches free-flying insects in wooded areas, fields, and over water, but also preys on insects on the water surface. Insects caught during flight are taken by swooping or dipping maneuvers. Most activity over water occurs between 1 to 2 m over the surface and the insects are taken by the mouth. Most feeding activity occurs about two hours after dark. (Fenton and Bell, 1979)

Little brown bats feed largely on aquatic insects. Midges are the primary source of food of M. lucifugus, but a large part of their diet comes from other aquatic insects. When available, beetles are easily identified by echolocation and easily captured. Other insects consumed include caddisflies, moths, mayflies, lacewings, and occasionally mosquitoes. (Anthony and Kunz, 1977; Fenton and Barclay, 1980)

  • Animal Foods
  • insects

Predation

Domestic cats have become adept at catching bats due to the close proximities of roosts to human habitations. Many predators take advantage of the high concentrations of bats in roosts. Predators such as martens and fishers take advantage of weak young that fall or hibernating individuals that are dislodged by grooming activities. Other predators of M. lucifugus include mice, owls, weasels, hawks, snakes, raccoons, domestic cats, and other small carnivores. (Fenton and Barclay, 1980; Griffin, 1958)

  • Anti-predator Adaptations
  • cryptic

Ecosystem Roles

Little brown bats have a major impact on the insect populations around their roosts. Active bats eat half of their body weight per night and lactating females eat more than their body weight per night. One M. lucifugus consumes approximately 3 to 7 grams of insects each night. (Barbour and Davis, 1969; Fenton and Barclay, 1980; Wilson and Ruff, 1999)

Tapeworms, and ectoparasites such as fleas, mites and bed bugs are carried by little brown bats. (Barbour and Davis, 1969; Fenton and Barclay, 1980; Fenton and Bell, 1979)

Commensal/Parasitic Species

Economic Importance for Humans: Positive

Members of this species are heavily researched and provide scientists with a bat model to test and study many aspects of the order, including echolocation, social behavior, feeding, and habitat use. Additionally, little brown bats eat pests that transmit diseases and eat agricultural products. They are also predators of mosquitoes and other pest around human habitats. (Barbour and Davis, 1969; Wilson and Ruff, 1999)

  • Positive Impacts
  • research and education
  • controls pest population

Economic Importance for Humans: Negative

Myotis lucifugus is the target of control measures due to the abundance of the species. These bats inhabit attics, roofs, trees, and other areas in close proximity to humans; therefore, homeowners have spent large amounts of money trying to erradicate M. lucifugus from these areas. Rabies transmission to humans is extremely low, and only a small percentage of M. lucifugus are infected with the disease. Although rabies in M. lucifugus is low, other parasites such as tapeworms, fleas, mites and bed bugs are common. (Fenton and Barclay, 1980)

Conservation Status

Myotis lucifugus is under no special conservation status as the species is abundant across North America. These bats thrives with expansion of human populations, as many of their roosting sites are built by humans. In spite of their overall abundance, some populations have suffered declines due to control measures and build-up of fat-soluble pesticides in their bodies. (Fenton and Barclay, 1980; Kunz, et al., 1977)

Temperate North American bats are now threatened by a fungal disease called “white-nose syndrome.” This disease has devastated eastern North American bat populations at hibernation sites since 2007. The fungus, Geomyces destructans, grows best in cold, humid conditions that are typical of many bat hibernacula. The fungus grows on, and in some cases invades, the bodies of hibernating bats and seems to result in disturbance from hibernation, causing a debilitating loss of important metabolic resources and mass deaths. Mortality rates at some hibernation sites have been as high as 90%. (Cryan, 2010; National Park Service, Wildlife Health Center, 2010)

Other Comments

There are six subspecies of M. lucifugus: M. l. alascensis, M. l. carissima, M. l. lucifugus, M. l. occultus, M. l. pernox, and M. l. relictus. Several of these subspecies were previously considered separate species: M. l. occultus, M. l. pernox, and M. l. carissima. (Fenton and Barclay, 1980; Hall, 1981)

Contributors

Tanya Dewey (editor), Animal Diversity Web.

Nancy Shefferly (editor), Animal Diversity Web.

Aaron Havens (author), University of Michigan-Ann Arbor, Phil Myers (editor, instructor), Museum of Zoology, University of Michigan-Ann Arbor.

Glossary

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.

World Map

acoustic

uses sound to communicate

agricultural

living in landscapes dominated by human agriculture.

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.

arboreal

Referring to an animal that lives in trees; tree-climbing.

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.

carnivore

an animal that mainly eats meat

causes disease in humans

an animal which directly causes disease in humans. For example, diseases caused by infection of filarial nematodes (elephantiasis and river blindness).

causes or carries domestic animal disease

either directly causes, or indirectly transmits, a disease to a domestic animal

chaparral

Found in coastal areas between 30 and 40 degrees latitude, in areas with a Mediterranean climate. Vegetation is dominated by stands of dense, spiny shrubs with tough (hard or waxy) evergreen leaves. May be maintained by periodic fire. In South America it includes the scrub ecotone between forest and paramo.

chemical

uses smells or other chemicals to communicate

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.

crepuscular

active at dawn and dusk

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.

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.

desert or dunes

in deserts low (less than 30 cm per year) and unpredictable rainfall results in landscapes dominated by plants and animals adapted to aridity. Vegetation is typically sparse, though spectacular blooms may occur following rain. Deserts can be cold or warm and daily temperates typically fluctuate. In dune areas vegetation is also sparse and conditions are dry. This is because sand does not hold water well so little is available to plants. In dunes near seas and oceans this is compounded by the influence of salt in the air and soil. Salt limits the ability of plants to take up water through their roots.

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.

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.

fertilization

union of egg and spermatozoan

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

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.

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.

insectivore

An animal that eats mainly insects or spiders.

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).

motile

having the capacity to move from one place to another.

mountains

This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.

native range

the area in which the animal is naturally found, the region in which it is endemic.

nocturnal

active during the night

pheromones

chemicals released into air or water that are detected by and responded to by other animals of the same species

polygynandrous

the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.

riparian

Referring to something living or located adjacent to a waterbody (usually, but not always, a river or stream).

seasonal breeding

breeding is confined to a particular season

sedentary

remains in the same area

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

social

associates with others of its species; forms social groups.

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.

suburban

living in residential areas on the outskirts of large cities or towns.

tactile

uses touch to communicate

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.

tropical savanna and grassland

A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.

savanna

A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.

temperate grassland

A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.

ultrasound

uses sound above the range of human hearing for either navigation or communication or both

urban

living in cities and large towns, landscapes dominated by human structures and activity.

visual

uses sight to communicate

viviparous

reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.

References

Anthony, E., T. Kunz. 1977. Feeding Strategies of the Little Brown Bat, Myotis Lucifugus, In Southern New Hampshire. Ecology, 58: 775-786.

Barbour, R., W. Davis. 1969. Bats of America. Lexington, Kentucky: The University Press of Kentucky.

Bassett, J., J. Wiebers. 1979. Subspecific Differences in the Urine Concentrating Ability of Myotis lucifugus. Journal of Mammalogy, 60(2): 395-397.

Belwood, J., M. Fenton. 1976. Variation in the diet of Myotis lucifugus (Chiroptera:Vespertilionidae). Canadian Journal of Zoology, 54: 1674-1678.

Cockrum, E. 1956. Homing, movements and longevity of bats. J. Mammal, 37: 48-57.

Cryan, P. 2010. "White-nose syndrome threatens the survival of hibernating bats in North America" (On-line). U.S. Geological Survey, Fort Collins Science Center. Accessed September 16, 2010 at http://www.fort.usgs.gov/WNS/.

Fenton, M., R. Barclay. 1980. Myotis lucifugus. Mammalian Species, 142: 1-8.

Fenton, M., G. Bell. 1979. Echolocation and feeding behaviour in four species of g.Myotis (Chiroptera). Canadian Journal of Zoology, 57: 1271-1277.

Griffin, D. 1958. Listening in the Dark. New Haven, Connecticut: Yale University Press.

Hall, E. 1981. The Mammals of North America. New York: John Wiley and Sons.

Koopman, K., F. Gudmundsson. 1966. American Museum Novitates. New York: American Museum of Natural History.

Kunz, T., E. Anthony, W. Rumage III. 1977. Mortality of little brown bats. J. Wildl. Manage, 41: 476-483.

Kurta, A. 1995. Mammals of the Great Lakes Region. Ann Arbor, MI: The University of Michigan Press.

Kurta, A., T. Kunz. 1988. Roosting Metabolic Rate and Body Tempature of Male Little Brown Bats (Myotis lucifugus) in Summer. Journal of Mammalogy, 69(3): 645-651.

National Park Service, Wildlife Health Center, 2010. "White-nose syndrome" (On-line). National Park Service, Wildlife Health. Accessed September 16, 2010 at http://www.nature.nps.gov/biology/wildlifehealth/White_Nose_Syndrome.cfm.

Nowak, R. 1994. Walker's Bats of the World. Baltimore, Maryland: The Johns Hopkins University Press.

Ratcliffe, J., J. Dawson. 2003. Behavioural flexibility: the little brown bat, Myotis lucifugus, and the northern long-eared but, M. septentrionalis, both glean and hawk prey. Animal Behaviour, 66: 847-856.

Schowalter, D. 1980. Swarming, Reproduction, and Early Hibernation of Myotis lucifugus and M. volans in Alberta, Canada. Journal of Mammalogy, 61(2): 350-354.

Tuttle, M. 1991. How North America's Bats Survive the Winter. Bats, 9(3): 7-12.

Wai-Ping, V., M. Fenton. 1988. Nonselective Mating in Little Brown Bats (Myotis lucifugus). Journal of Mammalogy, 69(3): 641-645.

Wilson, D., S. Ruff. 1999. The Smithsonian book of North American mammals. Washington D.C.: Smithsonian Institution Press in association with the American Society of Mammalogists.