Animal Diversity Web

Ge­o­graphic Range

Sil­ver-haired bats (La­sionyc­teris noc­ti­va­gans) are found in most of the United States, with the ex­cep­tion of the south­east and south­west­ern coasts of the United States. The species is found as far south as the Vic­to­ria province of Mex­ico. They in­habit areas as far north as the en­tire lower third of Canada, with a north­ward penin­su­lar ex­ten­sion fol­low­ing the coast into the lower south-cen­tral part of Alaska. Oc­ca­sion­ally, these bats have been doc­u­mented in Bermuda.

Dur­ing the sum­mer months, some pop­u­la­tions are sex­u­ally seg­re­gated, and it ap­pears that dis­tance and di­rec­tion of mi­gra­tion dif­fers among west­ern and east­ern pop­u­la­tions. It ap­pears that fe­males will con­sis­tently mi­grate greater dis­tances from hi­ber­nac­ula and spend the sum­mer fur­ther north­ward than males. West­ern pop­u­la­tions ap­pear to leave hi­ber­nac­ula and mi­grate east­ward and north­ward, while east­ern pop­u­la­tions only move north­ward. Pop­u­la­tions in British Co­lum­bia may not mi­grate, as equal sex ra­tios are noted through­out the year. Fe­males have been noted to be ab­sent in moun­tain­ous habi­tats in the west­ern U.S. dur­ing sum­mer months.

The only time the sexes are com­min­gled is dur­ing mi­gra­tion and hi­ber­na­tion. (Ar­royo-Cabrales, et al., 2008; Cryan, 2003; Kunz, 1982)

• Biogeographic Regions
• nearctic
  • native

Habi­tat

Sil­ver-haired bats are most com­monly found in bo­real or conif­er­ous and de­cid­u­ous for­est near bod­ies of water, such as rivers, lakes, streams, es­tu­ar­ies or ponds. Sum­mer day roosts are typ­i­cally under loose bark in trees, wil­lows (Salix), maple (Acer), and ash (Frax­i­nus) trees. These species may be most suit­able re­gard­ing the struc­ture of the bark and cav­i­ties they pro­vide (Kunz 1982a). They can also be found in dead trees, also called snags, that have loose bark to pro­vide cover, and cav­i­ties in these trees. These cav­i­ties are the lo­ca­tions of ma­ter­nity colonies, as well as sum­mer lo­ca­tions for bach­e­lor males. Un­com­monly, they use human struc­tures (garages, sheds, etc.) but roost­ing here tends to be soli­tary. Dur­ing the win­ter, these bats have been found in caves and other rocky area that pro­vide shel­ter, in tree cav­i­ties, and in build­ings. Win­ter finds tend to be anec­do­tal, as they don't seem to hi­ber­nate in large groups. (Ar­royo-Cabrales, et al., 2008; Kunz, 1982; Pa­triquin and Bar­clay, 2003; Perry, et al., 2010)

• Habitat Regions
• temperate
• terrestrial

• Terrestrial Biomes
• taiga
• forest
• scrub forest

• Aquatic Biomes
• rivers and streams

• Other Habitat Features
• suburban
• riparian
• caves

Phys­i­cal De­scrip­tion

The com­mon name for sil­ver-haired bats is de­rived from the sil­ver or white tips of the black fur that cov­ers both ven­tral and dor­sal sides of the an­i­mal. Their fur cov­ers al­most all of these bats ex­cept for the wings, snout, ears and pos­te­rior por­tion of the uropatag­ium mem­brane, which is a mem­brane that goes be­tween the bats legs that at­taches at the ankle. The wing and uropatag­ium mem­branes are dark brown to black. The ears are 5-9 mil­lime­ters and rounded at the tops. They have a slight up­turn to their snout. Adult males and fe­males look alike.

These bats as adults will weight be­tween 8 and 11 grams with the av­er­age being 9.5 grams. Their over­all length is 9.5 to 11.5 cen­time­ters with the av­er­age being 10 cen­time­ters. Their wingspan is 27 to 31 cen­time­ters with av­er­age being 29.5 cen­time­ters. Their right fore­arm, 3.7 to 4.4 cen­time­ters with the av­er­age being 3.9 cen­time­ters. Their basal meta­bolic rate is 3.73 to 4.66 cen­time­ters cubed of oxy­gen used per hour with the av­er­age being 4.14 cen­time­ters cubed of oxy­gen used per hour.

These bats' young, called pups, are born blind, with ears folded, ini­tially deaf. They will be born with no fur, but they will have dark skin in lo­ca­tions where fur will be found later as they de­velop. They will weigh 30 to 36% of the mother's body mass, which will about 2.0 g (range 1.4 to 2.3 g), de­pend­ing on how many young they pro­duce. (Kunz, 1982)

• Other Physical Features
• endothermic
• heterothermic
• bilateral symmetry

• Sexual Dimorphism
• sexes alike

• Range mass

  8 to 11 g

  0.28 to 0.39 oz

• Range length

  9.2 to 11.5 cm

  3.62 to 4.53 in

• Average length

  10 cm

  3.94 in

• Range wingspan

  27 to 31 cm

  10.63 to 12.20 in

• Average wingspan

  29.5 cm

  11.61 in

• Range basal metabolic rate

  3.79 to 4.66 cm3.O2/g/hr

• Average basal metabolic rate

  4.14 cm3.O2/g/hr

Re­pro­duc­tion

The males and fe­males are polyg­y­nan­drous and will form a swarm dur­ing mi­gra­tion. The sexes typ­i­cally do not com­min­gle dur­ing most of the year. Dur­ing this mi­gra­tion time, mat­ing oc­curs, and fe­males store the sperm until April or May. Then ges­ta­tion will start. After a ges­ta­tion pe­riod of 50 to 60 days, the fe­males will give live birth to typ­i­cally 2 pups. (Kunz, 1982; Van Schaik, et al., 2015)

• Mating System
• polygynandrous (promiscuous)

Sil­ver-haired bats breed once a year in the fall dur­ing mi­gra­tion. Fe­males will store sperm after mat­ing until the spring. After a ges­ta­tion pe­riod of 57 days, rang­ing from 50-60 days. The fe­male will give birth to (one to) two pups. The pups will weigh 1.4 to 2.3 grams with 2.0 grams being av­er­age. The young will nurse for ap­prox­i­mately 36 days (range 34 to 39) at which time they will be in­de­pen­dent. Age of sex­ual ma­tu­rity will range be­tween 145 to 160 days, with the av­er­age being 152 for both males and fe­males. (Kunz, 1982; Lacki, et al., 2007a)

• Key Reproductive Features
• iteroparous
• seasonal breeding
• gonochoric/gonochoristic/dioecious (sexes separate)
• sexual
• viviparous
• sperm-storing
• delayed fertilization

• Breeding interval

  Silver-haired bats breed once yearly.

• Breeding season

  Silver-haired bats breed in the fall.

• Range number of offspring

  1 to 2

• Average number of offspring

  2

• Average number of offspring

  2

  AnAge

• Range gestation period

  50 to 60 days

• Average gestation period

  57 days

• Range weaning age

  34 to 39 days

• Average weaning age

  36 days

• Range time to independence

  34 to 39 days

• Average time to independence

  36 days

• Range age at sexual or reproductive maturity (female)

  145 to 160 days

• Average age at sexual or reproductive maturity (female)

  152 days

• Range age at sexual or reproductive maturity (male)

  145 to 160 days

• Average age at sexual or reproductive maturity (male)

  152 days

Males have no in­vest­ment be­yond mat­ing in the fall. Fe­males are re­spon­si­ble for the pup or pups. Fe­males will nurse the young for an av­er­age of 36 days. Dur­ing this time the fe­male will for­age nor­mally. Prior to any kind of in­de­pen­dence, the pups re­main at­tached to the mother. The pups are born with teeth, so once in­de­pen­dent, they can fully for­age on their own. (Kunz, 1982)

• Parental Investment
• female parental care
• pre-hatching/birth
  • provisioning
    • female
• pre-weaning/fledging
  • provisioning
    • female
• pre-independence
  • provisioning
    • female

Lifes­pan/Longevity

Den­tal records have shown that sil­ver-haired bats can live as long as 12 years. Ex­pected lifes­pans in the wild have been re­ported as 7 years (range 5 to 10). These bats are not kept in cap­tiv­ity. (Kunz, 1982)

• Range lifespan
  Status: wild

  4 to 12 years

• Typical lifespan
  Status: wild

  5 to 10 years

• Average lifespan
  Status: wild

  7 years

Be­hav­ior

Sil­ver-haired bats are soli­tary an­i­mals that will seek shel­ter and sleep under loose bark, in dead trees or snags, in­side hol­low cav­i­ties of trees where heart rot may have taken place and holes that may have been used for birds and squir­rels. Very rarely will they seek shel­ter in­side homes. These bats are most com­monly found in wil­low, maple and ash trees. This is be­lieved to be due to the deep fis­sures in the bark. They will for­age mul­ti­ple times dur­ing the early evening through out the night. Typ­i­cal for­ag­ing times are early evening shortly after dark or right be­fore, and early evening times just be­fore dawn. Ac­cord­ing to Cryan (2003), the sep­a­ra­tion of times is due to avoid­ing fly­ing at the same times as red (La­si­u­rus bo­re­alis), hoary (La­si­u­rus cinereus), and big brown (Eptesi­cus fus­cus) bats. The red, hoary and big brown bats are faster fliers than the sil­ver-haired bats. Sil­ver-haired bats have been known to reach speeds of just 4.8-5.0 m/s; in order to be able to feed, they may wait for the oth­ers to be less ac­tive in suit­able habi­tats.

Dur­ing the sum­mer months sil­ver-haired males and fe­males are ge­o­graph­i­cally sep­a­rated from each other and will not be near one an­other until fall, when mi­gra­tion oc­curs. As they mi­grate for hi­ber­na­tion, they form swarms, at which time the two sexes meet. It is dur­ing this fall swarm that the fe­males and the males will mate, but the fe­males will not be­come preg­nant until after hi­ber­na­tion.

Hi­ber­na­tion of these bats will tend to hap­pen in more xeric, dry and warm, site dur­ing the win­ter. They will hi­ber­nate from Oc­to­ber until March.

These bats will use echolo­ca­tion as a form of find­ing food as well as a way to com­mu­ni­cate among bats. With wind tur­bines be­com­ing more preva­lent, echolo­ca­tion is not able to help them avoid the blades or the wind "vac­u­ums" that can kill them in­stantly. ("A Guide to the Bats of Vir­ginia", 2015; Cryan, 2003; Kunz, 1982; Lacki, et al., 2007b; Van Schaik, et al., 2015)

• Key Behaviors
• arboreal
• flies
• nocturnal
• motile
• migratory
• hibernation
• solitary

• Range territory size

  14 to 37 km^2

• Average territory size

  30 km^2

Home Range

Home range will varies for these bats, but will typ­i­cally in­clude a noc­tur­nal for­ag­ing area of about 30 square kilo­me­ters (range 14 to 37 km^2). They do not ac­tively de­fend a ter­ri­tory. (Kunz, 1982; Lacki, et al., 2007b)

Com­mu­ni­ca­tion and Per­cep­tion

Like most bats, sil­ver-haired bats use echolo­ca­tion, not only to per­ceive their en­vi­ron­ment but also as a way to hunt prey dur­ing the night. Ac­cord­ing to Voigt-Heucke et al. (2010), these bats use echolo­ca­tion as a way to iden­tify fa­mil­iar and un­fa­mil­iar in­di­vid­u­als. New­born pups iden­tify their moth­ers by scent. Con­trary to the phrase "blind as a bat," most bats can see as well as hu­mans dur­ing day­time hours.

Ac­cord­ing to Sil­vis (2016), who ex­am­ined 15 known acoustic files for sil­ver-haired bats, the av­er­age fre­quency at which these bats called was 29.26 kHz. The max­i­mum fre­quency of a call recorded for these bats was at 35.77 kHz, with the low­est at 26.39 kHz. The only species to be mea­sured with a lower fre­quency of bats in the mid-At­lantic U.S. is the hoary bat (La­si­urius cinereus).

These bats are gen­er­ally soli­tary, but do use vi­sual tac­tile com­mu­ni­ca­tion while mat­ing in the fall months. ("Clas­si­fi­ca­tion trees for acoustic iden­ti­fi­ca­tion of bat species in the mid-At­lantic", 2016; Voigt-Heucke, et al., 2010)

• Communication Channels
• visual
• tactile
• acoustic
• chemical

• Perception Channels
• visual
• tactile
• acoustic
• echolocation
• chemical

Food Habits

Ac­cord­ing to Lacki et al. (2007), sil­ver-haired bats will prey upon dif­fer­ent 11 dif­fer­ent or­ders of in­sects. In this study, moths in Order Lep­i­doptera made up over 40% of their diet. In many parts of their range, they are con­sid­ered moth spe­cial­ists. Flies and bee­tles made up most of the rest of their diet in this study. Other stud­ies sug­gested that they are op­por­tunists and that diet is re­liant on ge­o­graphic lo­ca­tion, time of the year and avail­abil­ity of prey. For ex­am­ple, the stom­ach con­tents of two bats were made up en­tirely (or nearly so) of cad­dis­flies (Tri­choptera). Spi­ders and ter­mites also have been found. These bats have been caught by mouse traps, and some re­searchers sug­gest that they for­age on the ground or glean. Fur­ther ev­i­dence of glean­ing are re­ports of sil­ver-haired bats tak­ing lar­vae di­rectly from trees. Data on food of these bats is de­rived from gut con­tent, di­rect ob­ser­va­tion of feed­ing, culled parts and most com­monly from feces or guano. Di­rect ob­ser­va­tion is rare. (Ar­royo-Cabrales, et al., 2008; Kunz, 1982; Lacki, et al., 2007b)

• Primary Diet
• carnivore
  • insectivore

• Animal Foods
• insects
• terrestrial non-insect arthropods

Pre­da­tion

The main preda­tors of sil­ver-haired bats are owls, such as the great horned owl Bubo vir­gini­anus. Mam­malian preda­tors in­clude striped skunks Mephi­tis mephi­tis that find the bats dur­ing times where they are not ac­tive. One case re­ported a rabid hoary bat La­si­u­rus cinereus killing a sil­ver-haired bat, but this was likely an iso­lated in­ci­dent. Re­cently, sharp-shinned hawks (Ac­cip­iter stria­tus) have be­come a major preda­tor be­cause of habi­tat changes. Log­ging has dis­rupted some of these bats' for­merly con­tigu­ous habi­tat, cre­at­ing frag­mented patches of suit­able habi­tat. This has caused a change in both feed­ing habits and time which they feed. These bats now nav­i­gate in more echolo­ca­tion clut­ter while for­ag­ing. The edge habi­tat in­cluded more open areas, which have made the bats more vul­ner­a­ble to preda­tors of owls and noc­tur­nal hawks. (Kunz, 1982; Pa­triquin and Bar­clay, 2003; Perry, et al., 2010)

• Anti-predator Adaptations
• cryptic

• Known Predators

  • great horned owls (Bubo virginianus)
  • sharp-shinned hawks (Accipiter striatus)
  • striped skunks (Mephitis mephitis)

Ecosys­tem Roles

Ac­cord­ing to Kunz (1982a), par­a­sites of sil­ver-haired bats in­clude mites in the fam­i­lies Sar­cop­ti­dae (Sar­cop­ti­dae la­sionyc­teris) and Lae­lap­i­dae (Lae­lap­i­dae my­otis). Most of the mites were found to be on the wing mem­brane or on top of the ears or in­side the ears. They also suf­fer from par­a­sites known as bat flies, which are wing­less and blind, and feed much like a tick. Two species of of bat flies from the fam­i­lies Nyc­teribi­idae (Basilia for­ci­pata) and Stre­bl­i­dae (Tri­chobius di­phyl­lae) have been doc­u­mented as using the sil­ver-haired bats as host. Bat bugs from the fam­ily Cimi­ci­dae (Cimex piosel­lus) are known par­a­sites. They've also been re­ported to host one flea species.

En­dopar­a­sites that have been found are ne­ma­todes (Cap­il­laria palmata), trema­todes (Acan­tha­trium lu­na­tum), and ces­todes (Hy­menolepis roud­abushi).

While there are many par­a­sites that use sil­ver-haired bats as a host, the par­a­sites found on or in a bat that lives in the east­ern part of the U.S. can often vary from species than those found on a sil­ver-haired bat who live in the west­ern part of the U.S. (Blanke­spoor and Ulmer, 1970; Boyd and Bern­stein, 1950; Kunz, 1982; Whitaker Jr. and East­erla, 1974)

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

Sil­ver-haired bat eco­nomic im­por­tance alone hasn’t been stud­ied, but the eco­nomic value of bats in the United States (as a whole) has been es­ti­mated by Boyles et al. (2011) to be $3.7 to 53 bil­lion per year as a nat­ural pest con­trol for crops. The bats also carry a species-spe­cific strain of ra­bies they are hence a focal an­i­mal for ra­bies re­search. (Boyles, et al., 2011; Mes­sen­ger, et al., 2002)

• Positive Impacts
• research and education
• controls pest population

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

Sil­ver-haired bats have been known to carry a species-spe­cific form of ra­bies virus that is known to in­fect hu­mans. Most ra­bies recordd are based on state health de­part­ment re­ports, and less than 5% of all bats sub­mit­ted for test­ing are sil­ver-haired bats. Those cases that can be at­trib­uted to sil­ver-haired bats can be done so via mol­e­c­u­lar test­ing of the in­fected per­son. The high­est con­cen­tra­tion of ra­bies cases are in the north­east­ern United States, be­cause that is where they are most com­mon dur­ing sum­mer months and most likely to in­ter­act with hu­mans. (Mes­sen­ger, et al., 2003; Mes­sen­ger, et al., 2002)

• Negative Impacts
• injures humans
  • carries human disease

Con­ser­va­tion Sta­tus

Sil­ver-haired bats are listed under as a species of "Least Con­cern" on list by the IUCN Red List, CITES and U.S. Fed­eral En­dan­gered List and in the state of Michi­gan. Ac­cord­ing to Adams and Ped­er­sen (2013), an in­crease of wind tur­bines being con­structed in mi­gra­tion cor­ri­dors, near or at for­ag­ing sites, or gen­er­ally within the sum­mer home ranges of in­di­vid­u­als, causes de­clines in pop­u­la­tions of sil­ver-haired bats. Log­ging also is a threat to their habi­tat con­ti­nu­ity. They are pro­posed as a Tier IV species in Vir­ginia, which means that they are cur­rently se­cure, but mul­ti­ple loom­ing threats (log­ging and wind tur­bines) causes con­cern for the species. In Mex­ico, where they are con­sid­ered a rare find, they are "sub­ject to spe­cial pro­tec­tion" (Ar­royo-Cabrales et al., 2008). ("A Guide to the Bats of Vir­ginia", 2015; Adams and Ped­er­sen, 2013; Ar­royo-Cabrales, et al., 2008)

• IUCN Red List

  Least Concern
  More information

• IUCN Red List

  Least Concern
  More information

• US Federal List

  No special status

• CITES

  No special status

• State of Michigan List

  No special status

Con­trib­u­tors

Justin Bent­ley (au­thor), Rad­ford Uni­ver­sity, Karen Pow­ers (ed­i­tor), Rad­ford Uni­ver­sity, Alex At­wood (ed­i­tor), Rad­ford Uni­ver­sity, Marisa Dameron (ed­i­tor), Rad­ford Uni­ver­sity, Tanya Dewey (ed­i­tor), Uni­ver­sity of Michi­gan-Ann Arbor.

Ref­er­ences

Vir­ginia De­part­ment of Game and In­land Fish­eries. A Guide to the Bats of Vir­ginia. Spe­cial Pub­li­ca­tion num­ber 5. Rich­mond, Vir­ginia: Vir­ginia De­part­ment of Game and In­land Fish­eries. 2015.

United States Ge­o­log­i­cal Sur­vey. Clas­si­fi­ca­tion trees for acoustic iden­ti­fi­ca­tion of bat species in the mid-At­lantic. none. Blacks­burg, Vir­ginia: United States Ge­o­log­i­cal Sur­vey. 2016.

Adams, R., S. Ped­er­sen. 2013. Bat Evo­lu­tion, Ecol­ogy, and Con­ser­va­tion. New York, New York: Springer Sci­ence & Busi­ness Media.

Ar­royo-Cabrales, J., B. Miller, F. Reid, A. Cuarón, P. de Gram­mont. 2008. "The IUCN Red List of Threat­ened Species 2008: e.T11339A3269157." (On-line). La­sionyc­teris noc­ti­va­gans. Ac­cessed Sep­tem­ber 15, 2015 at http://​www.​iucnredlist.​org/​details/​11339/​0.

Blanke­spoor, H., M. Ulmer. 1970. Helminths from six species of Iowa bats. Pro­ceed­ings of the Iowa Acad­emy of Sci­ence, 77: 200-206.

Boyd, E., M. Bern­stein. 1950. A new species of sar­cop­tic mite from a bat. Pro­ceed­ings of the En­to­mo­log­i­cal So­ci­ety of Wash­ing­ton, 52: 95–99.

Boyles, J., P. Cryan, G. Mc­Cracken, T. Kunz. 2011. Eco­nomic im­por­tance of bats in agri­cul­ture. Sci­ence, 332: 41-42.

Brack Jr, V., J. Whitaker Jr, S. Pruitt. 2004. Bats of Hoosier Na­tional For­est. Pro­ceed­ings of the In­di­ana Acad­emy of Sci­ence, 113/1: 76-87.

Cryan, P. 2003. Sea­sonal dis­tri­b­u­tion of mi­gra­tory tree bats (La­si­u­rus and La­sionyc­teris) in North Amer­ica. Jour­nal of Mam­mal­ogy, 84/2: 579-593.

Cryan, P. 2011. Wind tur­bines as land­scape im­ped­i­ments to the mi­gra­tory con­nec­tiv­ity of bats. En­vi­ron­men­tal Law, 41/355: 355-371.

Geluso, K., J. Hueb­schman, J. White, M. Bogan. 2004. Re­pro­duc­tion and sea­sonal ac­tiv­ity of sil­ver-haired bats (La­sionyc­teris noc­ti­va­gans) in west­ern Ne­braska. West­ern North Amer­i­can Nat­u­ral­ist, 64/3: 353-358.

Hock, R. 1951. The meta­bolic rates and body tem­pre­tures of bats. The Bi­o­log­i­cal Bul­letin, 101/3: 289-299.

Kalcou­nis, M., K. Hob­son, R. Bring­ham, K. Hecker. 1999. Bat ac­tiv­ity in the bo­real for­est: Im­por­tance of stand type and ver­ti­cal strata. Jour­nal of Mam­mal­ogy, 80/2: 673-682.

Kunz, T. 1982. Ecol­ogy of Bats. New York, New York and Lon­don, Eng­land: Pleneum Press.

Kunz, T. 1982. La­sionyc­teris noc­ti­va­gans. Mam­malian Species, 172: 1-5.

Lacki, M., L. Dodd, M. Baker. 2007. Prey con­sump­tion of in­sec­tiv­o­rous bats in conif­er­ous for­est of north-cen­tral Idaho. North­west Sci­ence, 81/3: 199-206.

Lacki, M., J. Hayes, A. Kurta. 2007. Bats in Forests. Bal­ti­more, Mary­land: Johns Hop­kins Uni­ver­sity Press.

McGuire, L., C. Guglielmo, S. Macken­zie, P. Tay­lor. 2012. Mi­gra­tory stopover in the long-dis­tance mi­grant sil­ver-haired bat, La­sionyc­teris noc­ti­va­gans. Jour­nal of An­i­mal Ecol­ogy, 81/2: 377–385.

Mes­sen­ger, S., J. Smith, L. Or­ciari, P. Yager, C. Rup­precht. 2003. Emerg­ing pat­tern of ra­bies deaths and in­creased viral in­fec­tiv­ity. Emerg­ing In­fec­tious Dis­eases, 9/2: 151-154.

Mes­sen­ger, S., J. Smith, C. Rup­precht. 2002. Emerg­ing epi­demi­ol­ogy of bat-as­so­ci­ated cryp­tic cases of ra­bies in hu­mans in the United States. Clin­i­cal In­fec­tious Dis­eases, 35/6: 738-747.

Misra, V., T. Du­mon­ceaux, J. Dubois, C. Willis, S. Nadin-Davis, A. Sev­erini, A. Wan­deler, R. Lind­say, H. Art­sob. 2009. De­tec­tion of poly­oma and corona viruses in bats of Canada. Jour­nal of Gen­eral Vi­rol­ogy, 90/8: 2015-2022.

Pa­triquin, K., R. Bar­clay. 2003. For­ag­ing by bats in cleared, thinned and un­har­vested bo­real for­est. Jour­nal of Ap­plied Ecol­ogy, 40/4: 646-657.

Perry, R., S. Carter, R. Thill. 2010. Tem­po­ral pat­terns in cap­ture rate and sex ratio of for­est bats in Arkansas. The Amer­i­can Mid­land Nat­u­ral­ist, 164/2: 270-282.

Van Schaik, J., R. Janssen, T. Bosch, A. Haarsma, J. Deker, B. Kranstauber. 2015. Bats swarm where they hi­ber­nate: Com­po­si­tional sim­i­lar­ity be­tween au­tumn swarm­ing and win­ter hi­ber­na­tion as­sem­blages at five un­der­ground sites. PLoS One, 10/7: 1-12.

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