Little brown bat
|Little brown bat|
(Le Conte, 1831)
The little brown bat (sometimes called little brown myotis) (Myotis lucifugus) is a species of mouse-eared bats found in North America. It has been a model organism for the study of bats (chiropterology).
|Relationships of Nearctic Myotis species|
It was described as a new species in 1831 by American naturalist John Eatton Le Conte. It was initially in the genus Vespertilio, with a binomial of Vespertilio lucifugus. "Lucifugus" is from Latin "lux" meaning "light" and "fugere" meaning "flee." The holotype had possibly been collected in Georgia near the Le Conte Plantation near Riceboro, but this has been disputed because the initial record lacked detail on where the specimen was collected.
Within its family, Vespertilionidae (the vesper bats), the little brown bat is a member of the subfamily Myotinae which contains only the mouse-eared bats of genus Myotis. Based on a 2007 study using mitochondrial and nuclear DNA, it is part of a Nearctic clade of mouse-eared bats. Its sister taxon is the Arizona myotis, M. occultus.
As of 2005, five subspecies of the little brown bat are recognized: M. l. lucifugus, M. l. alascensis, M. l. carissima, M. l. pernox, and M. l. relictus. Formerly, the Arizona myotis and southeastern myotis (M. austroriparius) were also considered subspecies (M. l. occultus and M. l. austroriparius), but both are now recognized as full species. In a 2018 study by Morales and Carstens, they concluded that the five subspecies are independent, paraphyletic lineages that each warrant specific status.
The little brown bat is a small species, with individuals weighing 5.5–12.5 g (0.19–0.44 oz) with a total body length of 8–9.5 cm (3.1–3.7 in). Individuals have the lowest weight in the spring as they emerge from hibernation. It has a forearm length of 36–40 mm (1.4–1.6 in) and a wingspan of 22.2–26.9 cm (8.7–10.6 in). It as a sexually dimorphic species, with females larger than males on average. A variety of fur colors is possible, with pelage ranging from pale tan or reddish to dark brown. Its belly fur is a lighter color than its dorsal fur. Its fur is glossy in appearance, though less so on its belly. A variety of pigmentation disorders have been documented in this species, including albinism (total lack of pigment), leucism (partial lack of pigment), and melanism (over-pigmentation).
Head and teeth
It is a diphyodont mammal, meaning that it has two sets of teeth during its lifetime—milk teeth and adult teeth. The dental formula of the milk teeth is 18.104.22.168 for a total of 22 teeth, while that of the adult teeth is 22.214.171.124 for a total of 38 teeth. Newborns ("pups") are born with 20 milk teeth which becomes 22 when the final upper premolars emerge. Pups begin losing milk teeth once they have reached a body length of 55–60 mm (2.2–2.4 in). Loss of all the milk teeth and emergence of all the permanent teeth is usually complete by the time a juvenile is 80 mm (3.1 in) long. This corresponds with gaining the ability to fly; juveniles can fly by 18 days old, which is when they begin consuming the adult diet of insects.
It has a relatively short snout and a gently sloped forehead. It lacks a sagittal crest, which can be used to distinguish it from the Arizona myotis. Its skull length is 14–16 mm (0.55–0.63 in). The braincase appears nearly circular though somewhat flattened when viewed from the back. Its ears are 11–15.5 mm (0.43–0.61 in) long, while the tragi are 7–9 mm (0.28–0.35 in) long. The tragi are blunt at the tips and considered of medium length for a mouse-eared bat.
The little brown bat is dichromatic and its eyesight is likely sensitive to ultraviolet and red light, based on a genetic analysis that discovered that the genes SWS1 and M/LWS were present and functional. Its ability to see ultraviolet light may be useful in capturing insects, as 80% of nocturnal moths' wings reflect UV light. It is unclear if or how seeing red light is advantageous for this species.
Relative to frugivorous bat species such as the Jamaican fruit bat, the little brown bat has small eyes and a reduced olfactory epithelium. Instead, it has a more sophisticated system of echolocation, suggesting that reliance on echolocation decreases the need for orientation via sight or smell.
The little brown bat can be confused with the Indiana bat (M. sodalis) in appearance. The two can be differentiated by the little brown bat's lack of a keeled calcar—the cartilaginous spur on its uropatagium is not as pronounced. Additionally, the little brown bat can be distinguished by the presence of hairs on its toes and feet that extend beyond the length of the digits. The northern long-eared bat (M. septentrionalis), another similar species, can be distinguished by its much longer ears and tragi that are long and sharply pointed.
Biology and ecology
Reproduction and life cycle
Little brown bats have a promiscuous mating structure, meaning that individual bats of both sexes mate with multiple partners. It is a seasonal breeder, with mating taking place in the fall before the annual hibernation. As a seasonal breeder, males do not produce sperm year-round; instead, spermatogenesis occurs May through August each year. Throughout the spring and summer, males and females roost separately. In the fall, however, individuals of both sexes will congregate in the same roost in a behavior known as "swarming," defined as "mass visitations by bats to underground sites prior to or just following hibernation." Like several other bat species, males of this species exhibit homosexual behaviors, with male bats mating indiscriminately with torpid, roosting bats, regardless of sex.
Although copulation occurs in the fall, fertilization does not occur until the spring due to sperm storage. Gestation proceeds for 50-60 days following fertilization. The litter size is one individual. At birth, pups weigh approximately 2.2 g (0.078 oz) and have a forearm length less than 17.2 mm (0.68 in). Pups' eyes and ears are closed at first, but open within a few hours of birth. They exhibit rapid growth; at around three weeks old, the young start flying, begin the weaning process, and are of a similar size to adults in forearm length but not weight. Females, but not males, may become sexually mature in the first year of life.
Little brown bats are very long-lived animals relative to their body sizes. In the wild, individuals have been documented living up to 34 years. Males and females have high annual survival rates (probability of surviving another year), though survival rates vary by sex and region. One colony documented in Ontario had a male survival rate of 81.6% and a female survival rate of 70.8%; a colony in southern Indiana had survival rates of 77.1% and 85.7% for males and females, respectively.
Diet and foraging
Little brown bats produce calls that are high intensity frequency modulated (FM) and that last from less than one millisecond (ms) to about 5 ms and have a sweep rate of 80–40 kHz, with most of their energy at 45 kHz. Bats usually emit 20 calls per second when in flight. When pursuing prey, a bat emits 200 calls per second. It also emits a high-pulse repetitive call if it wants to land.
Bats that are in danger of colliding will reduce the terminal portion of their sweep calls to 25 kHz, creating a "honking" sound. The bats also find roosting sites by listening to the echolocation calls of other individuals. Some complex vocalizations are used by mothers and their pups.
Little brown bats are insectivores, eating moths, wasps, beetles, gnats, mosquitoes, midges and mayflies, among others. Since many of their preferred meals are insects with an aquatic life stage, such as mosquitoes, they prefer to roost near water. Brown bats forage near bodies of water and move in and out of adjacent vegetation. Evening forages are done in groups and above the water. They echolocate to find their prey. They are particularly good at hunting insects when they are at close range and packed together. When hunting, little brown bats capture prey both by gleaning and by catching them in the air. When in flight, bats scoop up the prey with their wings, while prey above water is directly grabbed with the mouth.
Brown bats do not claim feeding areas like a territory, however individuals frequently return to the same feeding sites where they have previously made successful catches. When hunting swarms, brown bats usually select no more than two species. They feed on more species when they are scattered. If they do not catch any food, they will enter a torpor similar to hibernation that day, awakening at night to hunt again.
The bats' diet makes this species beneficial to agriculture as it eats many species of agricultural pests.
Predation and disease
The presence of helminth parasites in 'M lucifugus is most common in the spring and fall and least common in the summer. Digenetic trematodes are the most common of these parasites, with the more common of these species including Ototrema schildti and Plagiorchis vespertilionis.
Range and habitat
The little brown bat lives throughout much of North America. In the north, its range extends as far west as Alaska and across much of Canada to Labrador. In the south, its range extends to Southern California and across the northern parts of Arizona and New Mexico. Historically, the largest known aggregations of this species occurred in the karstic regions of the Eastern United States. Little brown bats have been found in Iceland, Alaska, and Kamchatka, likely due to accidental ship transportation by humans.
The little brown bat inhabits three different roosting sites: day roosts, night roosts, and hibernation roosts. Bats use day and night roosts during spring, summer, and fall, while hibernacula are used in winter. As with most bats, the little brown bat is mostly active at night and leaves its roost at dusk and the next two or three hours are peak activity periods. Since little brown bats live in a temperate zone, they must find some way of dealing with winter. Most temperate bats either migrate or hibernate, but little brown bats do both. In summer, the males and females live apart, and the females raise the young. When fall comes, both sexes fly south to a hibernaculum, where they mate and then hibernate. Little brown bats undergo a prolonged period of hibernation during the winter due to the lack of food. They hibernate in caves as a community.
Day roosts are usually in buildings or trees, under rocks or wood piles, and sometimes in caves. Nursery roosts are in natural hollows and buildings, or at least close to them. Little brown bats typically occupy darkly lit sites for day roosts, though open areas are occasionally used; some roosts have been found under the sheet metal roofs of trappers' caches and attics of buildings. The most important aspects of day roosts and nurseries are the ambient temperature and availability of shelter; warmth is especially important to ensure the growth of young bats. Hibernacula are typically not diurnally roosted in for this reason, as male bats freely expend energy outside of hibernation and thus avoid these low ambient temperatures and young bats require warmer ambient temperatures for growth, though some bats swarm near or in hibernacula. Water vapor levels and temperature in roosts effect rates of water loss, and most diurnal water loss is through the lungs and sweat glands. Adult males and nonparous females inhabit day roosts away from the nurseries, with these day roosts somewhat cooler than nurseries. Males tend to forage close to their day roosts, while females can forage upwards of 5 km (3.1 mi) or even 6 km (3.7 mi) away from day roosts.
Night roosts tend to be in the same buildings as day roosts, but these roosts tend to be in different spots that are more constrained; the bats pack together for warmth. Bats rest in night roosts after feeding in the evening which may serve to keep their feces away from the day roosts and thus less noticeable to predators. Further evidence for the importance of roost temperature is the little brown bat's tendency to rarely occupy night roosts when temperatures are above 59 °F (15 °C).
Little brown bats typically hibernate in caves or unused mines. High humidity and temperatures above freezing are common to hibernation sites, though there have been instances of little brown bats hibernating at sites below freezing. M. lucifugus prefers to hibernate at temperatures around 35.5 °F (1.9 °C), though it is less likely that the bats have an increased sense of temperature and more likely that the bats remain in torpor longest at 35.5 °F, meaning the increase in activity at temperatures outside of 35.5 °F makes the bats tend to roost towards 35.5 °F. During hibernation, levels of water loss are not correlated to body mass or ambient temperature but to differences in water vapor pressure in the bat's tissue and the atmosphere, though bats can lose more water when leaving winter torpor. Bats often leave the state of winter torpor due to increased urine waste levels, leaving the torpor state to urinate, defecate, and drink. Northern populations of bats enter hibernation in early September and end in mid-May, while southern populations enter in November and ends mid-March. Little brown bats are true hibernators.
As of 2018, the little brown bat's conservation status had not been reevaluated by the IUCN since 2008 when it was designated as the lowest conservation priority, least concern. Until recently, the species was regarded as one of the most common bats in North America. However, a serious threat to the species has emerged in the form of a fungus-caused disease known as white-nose syndrome. It was one of the first bat species documented with the disease, which now affects at least seven hibernating bat species in the United States and Canada. From 2005 to 2011, over one million little brown bats died from the disease in the Northeastern United States, with winter hibernacula populations declining up to 99%. As of 2017, hibernacula counts for little brown bats in the Northeast had declined by an average of 90%.
White-nose syndrome first appeared in New York in 2006; it has steadily diffused from eastern New York, though, until recently, remaining east of the Rocky Mountains. In March 2016, white-nose syndrome was detected on a little brown bat in King County, Washington, representing a 1,300 mi (2,100 km) jump from the previous westernmost extent of the disease in any bat species.
In 2010, Frick et al. predicted a 99% chance of local extinction of little brown bats by the year 2026. They also predicted that the pre-white-nose syndrome population of 6.5 million individuals could be reduced to as few as 65,000 (1%) via the disease outbreak. Despite heavy declines, the species has avoided extinction in the Northeast through the persistence of small, localized populations. While the mortality rate of the disease is very high, some individuals that are exposed do survive.
In 2010, Kunz and Reichard published a report arguing that the precipitous decline of the little brown bat justified its emergency listing as a federally endangered species under the U.S. Endangered Species Act. However, it is not federally listed as threatened or endangered as of 2018, though several U.S. states list it as endangered (Connecticut, Maine, Massachusetts, New Hampshire, Vermont, Virginia), threatened (Tennessee, Wisconsin), or of Special Concern (Michigan, Ohio)
The little brown bat was listed as an endangered species by the Committee on the Status of Endangered Wildlife in Canada in February 2012 after an emergency assessment. The emergency designation as endangered was confirmed in November 2013.
Relationship to people
Little brown bats commonly occupy human structures. Females will situate maternity colonies within buildings. This small body size of this species can make it challenging to prevent individuals from entering a structure, as they can take advantage of gaps or holes as small as 1.5 in (3.8 cm) x 0.25 in (0.64 cm). Once inside a building, a colony of little brown bats can disturb human inhabitants with their vocalizations and production of feces (guano) and urine. Large accumulations of guano can provide a growth medium for fungi, including the species that causes histoplasmosis. Concerns about humans becoming affected by bat ectoparasites such as ticks, fleas, or bat bugs are generally unfounded, as parasites that feed on bats are often specific to bats and die without them.
Because they are often found in proximity to humans, the little brown bat and the not-closely related big brown bat are the two bat species most frequently submitted for rabies testing in the United States. Little brown bats infrequently test positive for the rabies virus; of the 586 individuals submitted for testing across the United States in 2015, the most recent data available as of 2018, 16 (2.7%) tested positive for the virus.
Little brown bats are a species that will use bat houses for their roosts. Landowners will purchase or construct bat houses and install them, hoping to attract bats for various reasons. Some install bat houses in an attempt to negate the effects of removing a colony from a human structure ("rehoming" them into a more acceptable space). While this can be effective for other species, there is not evidence to suggest that this is effective for little brown bats. Others are attempting to help bats out of concern for them due to the effects of white-nose syndrome. Bat houses are also installed in an attempt to control the bats' insect prey such as mosquitoes or taxa that harm crops.
- Arroyo-Cabrales, J. & Álvarez-Castañeda, S.T. (2008). "Myotis lucifugus". The IUCN Red List of Threatened Species. 2008: e.T14176A4415629. doi:10.2305/IUCN.UK.2008.RLTS.T14176A4415629.en. Retrieved 27 December 2017.
- Stadelmann, B; Lin, L.K; Kunz, T.H; Ruedi, M (2007). "Molecular phylogeny of New World Myotis (Chiroptera, Vespertilionidae) inferred from mitochondrial and nuclear DNA genes". Molecular Phylogenetics and Evolution. 43 (1): 32–48. doi:10.1016/j.ympev.2006.06.019. PMID 17049280.
- Cuvier, Georges; Baird, Spencer Fullerton; Goode, G. Brown; Latreille, P. A.; Laurillard, Charles Léopold; Mason, Otis Tufton; McElfresh, Henry; McMurtrie, Henry; Schoolcraft, Henry Rowe (1831). "The animal kingdom arranged in conformity with its organization / by the Baron Cuvier ; the Crustacea, Arachnides and Insecta, by P.A. Latreille ; translated from the French, with notes and additions, by H. M'Murtrie ; in four volumes, with plates". 1. G. & C. & H. Carvill: 431. doi:10.5962/bhl.title.41463.
- Stangl, F. B.; Christiansen, P. G.; Galbraith, E. J. (1993). "Abbreviated guide to pronunciation and etymology of scientific names for North American land mammals north of Mexico" (PDF). Occasional Papers, the Museum, Texas Tech University (154): 7.
- Fenton, M. Brock; Barclay, Robert M. R. (1980). "Myotis lucifugus". Mammalian Species (142): 1–8. doi:10.2307/3503792. JSTOR 3503792.
- Davis, Wayne H.; Rippy, Charles L. (1968). "Distribution of Myotis lucifugus and Myotis austroriparius in the Southeastern United States". Journal of Mammalogy. 49 (1): 113–117. doi:10.2307/1377733. JSTOR 1377733.
- Hoofer, Steven R.; Bussche, Ronald A. Van Den (2003). "Molecular Phylogenetics of the Chiropteran Family Vespertilionidae". Acta Chiropterologica. 5: 1–63. doi:10.3161/001.005.s101.
- Wilson, D.E.; Reeder, D.M., eds. (2005). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. ISBN 978-0-8018-8221-0. OCLC 62265494.
- Piaggio, Antoinette J.; Valdez, Ernest W.; Bogan, Michael A.; Spicer, Greg S. (2002). "Systematics of Myotis occultus (Chiroptera: Vespertilionidae) Inferred from Sequences of Two Mitochondrial Genes". Journal of Mammalogy. 83 (2): 386–395. doi:10.1644/1545-1542(2002)083<0386:SOMOCV>2.0.CO;2.
- Jones, Clyde; Manning, Richard W. (1989). "Myotis austroriparius". Mammalian Species (332): 1–3. doi:10.2307/3504306. JSTOR 3504306.
- Morales, Ariadna E.; Carstens, Bryan C. (2018). "Evidence that Myotis lucifugus "Subspecies" are Five Nonsister Species, Despite Gene Flow". Systematic Biology. 67 (5): 756–769. doi:10.1093/sysbio/syy010. PMID 29462459.
- Wisconsin Department of Natural Resources (2013). Wisconsin Little Brown Bat Species Guidance (PDF) (Report). Madison, Wisconsin: Bureau of Natural Heritage Conservation, Wisconsin Department of Natural Resources. PUB-ER-705.
- Buchanan, G. Dale (1985). "Comments on Frequency of Melanism in Myotis lucifugus". Journal of Mammalogy. 66 (1): 178. doi:10.2307/1380979. JSTOR 1380979.
- Fenton, M. Brock (1970). "The deciduous dentition and its replacement in Myotis lucifugus (Chiroptera: Vespertilionidae)". Canadian Journal of Zoology. 48 (4): 817–820. doi:10.1139/z70-143.
- "Little Brown Myotis - Myotis lucifugus". Montana Field Guides. Montana Natural Heritage Program and Montana Fish, Wildlife and Parks. Retrieved 3 November 2018.
- Zhao, Huabin; Xu, Dong; Zhou, Yingying; Flanders, Jon; Zhang, Shuyi (2009). "Evolution of opsin genes reveals a functional role of vision in the echolocating little brown bat (Myotis lucifugus)". Biochemical Systematics and Ecology. 37 (3): 154–161. doi:10.1016/j.bse.2009.03.001.
- Bhatnagar, Kunwar P. (1975). "Olfaction in Artibeus jamaicensis and Myotis lucifugus in the context of vision and echolocation" (PDF). Experientia. 31 (7): 856. doi:10.1007/BF01938504. PMID 1140332.
- Thomas, Donald W.; Fenton, M. Brock; Barclay, Robert M. R. (1979). "Social Behavior of the Little Brown Bat, Myotis lucifugus: I. Mating Behavior" (PDF). Behavioral Ecology and Sociobiology. 6 (2): 129–136. doi:10.1007/BF00292559.
- Burns, Lynne E.; Frasier, Timothy R.; Broders, Hugh G. (2014). "Genetic connectivity among swarming sites in the wide ranging and recently declining little brown bat (Myotis lucifugus)". Ecology and Evolution. 4 (21): 4130–4149. doi:10.1002/ece3.1266. PMC 4242565. PMID 25505539.
- Riccucci, M. (2010). "Same-sex sexual behaviour in bats". Hystrix, the Italian Journal of Mammalogy. 22 (1). doi:10.4404/Hystrix-22.1-4478.
- Kunz, T. H.; Anthony, E. L. P. (1982). "Age Estimation and Post-Natal Growth in the Bat Myotis lucifugus". Journal of Mammalogy. 63: 23–32. doi:10.2307/1380667. JSTOR 1380667.
- Brunet-Rossinni, Anja K. (2004). "Reduced free-radical production and extreme longevity in the little brown bat (Myotis lucifugus) versus two non-flying mammals". Mechanisms of Ageing and Development. 125 (1): 11–20. doi:10.1016/j.mad.2003.09.003. PMID 14706233.
- Keen, R.; Hitchcock, H. B. (1980). "Survival and Longevity of the Little Brown Bat (Myotis lucifugus) in Southeastern Ontario". Journal of Mammalogy. 61: 1–7. doi:10.2307/1379951. JSTOR 1379951.
- Fenton, M. Brock; Bell, Gary P. (1979). "Echolocation and feeding behaviour in four species of Myotis (Chiroptera)" (PDF). Canadian Journal of Zoology. NRC Research Press. 57 (6): 1271–1277.
- Belwood, J. J.; Fenton, M. B. (1976). "Variation in the diet of Myotis lucifugus (Chiroptera: Vespertilionidae)". Canadian Journal of Zoology. NRC Research Press. 54 (10): 1674–1678.
- Ratcliffe, John M.; Dawson, Jeff W. (2003). "Behavioural flexibility: the little brown bat, Myotis lucifugus, and the northern long-eared bat, M. septentrionalis, both glean and hawk prey" (PDF). Animal Behaviour. Academic Press. 66 (5): 847–856.
- Kunz, Thomas H.; McCracken, Gary F.; Cryan, Paul M.; Boyles, Justin G. (2011). "Economic importance of bats in agriculture" (PDF). Science. American Association for the Advancement of Science. 332 (6025): 41–42.
- Coggins, James R.; Tedesco, John L.; Rupprecht, Charles E. (1982). "Seasonal changes and overwintering of parasites in the bat, Myotis lucifugus (Le Conte), in a Wisconsin hibernaculum" (PDF). American Midland Naturalist. JSTOR: 305–315.
- Vonhof, Maarten J.; Russell, Amy L.; Miller-Butterworth, Cassandra M. (2015). "Range-Wide Genetic Analysis of Little Brown Bat (Myotis lucifugus) Populations: Estimating the Risk of Spread of White-Nose Syndrome". PLOS ONE. 10 (7): e0128713. doi:10.1371/journal.pone.0128713. PMC 4495924. PMID 26154307.
- Youngman, P. M. (1975). Mammals of the Yukon Territory. National Museum of Natural Sciences, National Museums of Canada.
- Davis, Wayne H.; Hitchcock, Harold B. (1965). "Biology and migration of the bat, Myotis lucifugus, in New England" (PDF). Journal of Mammalogy. American Society of Mammalogists. 46 (2): 296–313.
- Kurta, Allen; Kunz, Thomas H. (1988). "Roosting metabolic rate and body temperature of male little brown bats (Myotis lucifugus) in summer" (PDF). Journal of Mammalogy. JSTOR. 69 (3): 645–651.
- Randall, Lea A.; Jung, Thomas S.; Barclay, Robert M.R. (2014). "Roost-site selection and movements of little brown myotis (Myotis lucifugus) in southwestern Yukon" (PDF). Northwestern Naturalist. BioOne. 95 (3): 312–317.
- McManus, John J. (1974). "Activity and thermal preference of the little brown bat, Myotis lucifugus, during hibernation" (PDF). Journal of Mammalogy. JSTOR. 55 (4): 844–846.
- Thomas, Donald W.; Cloutier, Danielle (1992). "Evaporative water loss by hibernating little brown bats, Myotis lucifugus" (PDF). Physiological Zoology. University of Chicago Press. 65 (2): 443–456.
- Macrae, Maria (2018). "Little Brown Bat". Hinterland Who's Who, Canadian Wildlife Federation. Retrieved 14 August 2017.
- Miller-Butterworth, C. M.; Vonhof, M. J.; Rosenstern, J.; Turner, G. G.; Russell, A. L. (2014). "Genetic Structure of Little Brown Bats (Myotis lucifugus) Corresponds with Spread of White-Nose Syndrome among Hibernacula". Journal of Heredity. 105 (3): 354–364. doi:10.1093/jhered/esu012. PMID 24591103.
- Dzal, Y.; McGuire, L. P.; Veselka, N.; Fenton, M. B. (2011). "Going, going, gone: The impact of white-nose syndrome on the summer activity of the little brown bat (Myotis lucifugus)". Biology Letters. 7 (3): 392–394. doi:10.1098/rsbl.2010.0859. PMC 3097845. PMID 21106570.
- Dobony, Christopher A.; Johnson, Joshua B. (2018). "Observed Resiliency of Little Brown Myotis to Long-Term White-Nose Syndrome Exposure". Journal of Fish and Wildlife Management. 9: 168–179. doi:10.3996/102017-JFWM-080.
- "White-nose Syndrome: A Deadly Disease". batcon.org. Bat Conservation International. Retrieved 4 November 2018.
- Frick, W. F.; Pollock, J. F.; Hicks, A. C.; Langwig, K. E.; Reynolds, D. S.; Turner, G. G.; Butchkoski, C. M.; Kunz, T. H. (2010). "An Emerging Disease Causes Regional Population Collapse of a Common North American Bat Species". Science. 329 (5992): 679–682. doi:10.1126/science.1188594. PMID 20689016.
- Kunz, TH; Reichard, JD. (2010). Status review of the little brown myotis (Myotis lucifugus) and determination that immediate listing under the Endangered Species Act is scientifically and legally warranted. Status Review prepared for US Fish and Wildlife Service (PDF) (Report).
- State of Connecticut Department of Energy and Environmental Protection Bureau of Natural Resources (2015). "Connecticut's Endangered, Threatened and Special Concern Species" (PDF). Connecticut.gov.
- "Myotis lucifugus (Little Brown Bat" (PDF). State of Maine. 13 January 2016.
- Natural Heritage & Endangered Species Program. "Little Brown Myotis Myotis lucifugus" (PDF). Massachusetts Division of Fisheries & Wildlife.
- "Endangered and Threatened Wildlife of NH". New Hampshire Fish and Game. New Hampshire Fish and Game Department. Retrieved 4 November 2018.
- "Little brown bat". Vermont Fish & Wildlife Department. State of Vermont. Retrieved 4 November 2018.
- Virginia Department of Game and Inland Fisheries. "Special Status Faunal Species in Virginia" (PDF). Virginia.gov. Retrieved 4 November 2018.
- "Rules and Regulation for In Need of Management, Threatened, and Endangered Species" (PDF). Tennessee.gov. Retrieved 4 November 2018.
- "Michigan's Rare Animals". Michigan Natural Features Inventory. Michigan State University. Retrieved 4 November 2018.
- Ohio Department of Natural Resources Division of Wildlife (July 2018). Ohio's Listed Species (PDF) (Report). R0718.
- COSEWIC (2013). COSEWIC assessment and status report on the Little Brown Myotis Myotis lucifugus, Northern Myotis Myotis septentrionalis and Tri-colored Bat Perimyotis subflavus in Canada (Report). Ottawa: Committee on the Status of Endangered Wildlife in Canada.
- Neilson, A.; Fenton, M. (1994). "Responses of Little Brown Myotis to Exclusion and to Bat Houses". Wildlife Society Bulletin (1973-2006). 22 (1): 8–14. JSTOR 3783215.
- Greenhall, Arthur M.; Frantz, Stephen C. (1994). "Bats". The Handbook: Prevention and Control of Wildlife Damage.
- Davis, A. D.; Jarvis, J. A.; Pouliott, C. E.; Morgan, S. M. D.; Rudd, R. J. (2013). "Susceptibility and Pathogenesis of Little Brown Bats (Myotis lucifugus) to Heterologous and Homologous Rabies Viruses". Journal of Virology. 87 (16): 9008–9015. doi:10.1128/JVI.03554-12. PMC 3754046. PMID 23741002.
- Birhane, Meseret G.; Cleaton, Julie M.; Monroe, Ben P.; Wadhwa, Ashutosh; Orciari, Lillian A.; Yager, Pamela; Blanton, Jesse; Velasco-Villa, Andres; Petersen, Brett W.; Wallace, Ryan M. (2017). "Rabies surveillance in the United States during 2015". Journal of the American Veterinary Medical Association. 250 (10): 1117–1130. doi:10.2460/javma.250.10.1117. PMID 28467751.
- Tuttle, Merlin; Hensley, Donna (1993). "Bat Houses: The Secrets of Success". batcon.org. Bat Conservation International. Retrieved 5 November 2018.
- "How You Can Help". White-Nose Syndrome Response Team. U.S. Fish and Wildlife Service, Department of the Interior. Retrieved 5 November 2018.
- Long, Rachael Freeman; Simpson, Tiffanie; Ding, Tzung-Su; Heydon, Steve; Reil, Wilbur (1998). "Bats feed on crop pests in Sacramento Valley". California Agriculture. 52: 8–10. doi:10.3733/ca.v052n01p8.