Corynorhinus townsendii

(Cooper, 1837)

Townsend's Big-eared Bat

G4Apparently Secure Found in 41 roadless areas NatureServe Explorer →
G4Apparently SecureGlobal Rank
Least concernIUCN
PSESA Status
MediumThreat Impact
Corynorhinus townsendii. Photo by Juan Cruzado Cortés, via iNaturalist.
Juan Cruzado Cortés, CC BY 4.0
Corynorhinus townsendii. Photo by Juan Cruzado Cortés, via iNaturalist.
Juan Cruzado Cortés, CC BY 4.0
Corynorhinus townsendii. Photo by Ian McFaul, via iNaturalist.
Ian McFaul, CC BY 4.0
Corynorhinus townsendii. Photo by Pinnacles National Park, via iNaturalist.
Pinnacles National Park, CC BY 4.0
Corynorhinus townsendii. Photo by Judi Sanders, via iNaturalist.
Judi Sanders, CC BY 4.0
Virginia big-eared bat (Corynorhinus (=Plecotus) townsendii virginianus). Photo by U.S. Fish & Wildlife Service, Public Domain (U.S. Government Work), via ECOS.
U.S. Fish & Wildlife Service, https://www.usa.gov/government-works
Ozark big-eared bat (Corynorhinus (=Plecotus) townsendii ingens). Photo by U.S. Fish & Wildlife Service, Public Domain (U.S. Government Work), via ECOS.
U.S. Fish & Wildlife Service, https://www.usa.gov/government-works
Identity
Unique IDELEMENT_GLOBAL.2.103228
Element CodeAMACC08010
Record TypeSPECIES
ClassificationSpecies
Classification StatusStandard
Name CategoryVertebrate Animal
IUCNLeast concern
Endemicoccurs (regularly, as a native taxon) in multiple nations
KingdomAnimalia
PhylumCraniata
ClassMammalia
OrderChiroptera
FamilyVespertilionidae
GenusCorynorhinus
USESAPS
Synonyms
Plecotus townsendiiCooper, 1837
Other Common Names
Oreillard de Townsend (FR) Townsend's big-eared bat (EN) Un Murciélago (ES)
Concept Reference
Baker, R. J., L. C. Bradley, R. D. Bradley, J. W. Dragoo, M. D. Engstrom, R. S. Hoffman, C. A. Jones, F. Reid, D. W. Rice, and C. Jones. 2003a. Revised checklist of North American mammals north of Mexico, 2003. Museum of Texas Tech University Occasional Papers 229:1-23.
Taxonomic Comments
This bat was included in the genus Plecotus by Handley (1959).
Conservation Status
Rank MethodLegacy Rank calculation - Excel v3.1x
Review Date2016-04-04
Change Date2015-03-16
Edition Date2015-03-25
Edition AuthorsHammerson, G.
Threat ImpactMedium
Range Extent>2,500,000 square km (greater than 1,000,000 square miles)
Rank Reasons
Large range spans North America; very large number of known roost sites and locations; substantial overall population size, but local populations tend to be small; overall recent trend not well known; apparently declining in abundance in portions of the range but stable or increasing in other areas; increasing in abundance in the east as a result of effective conservation measures; many roosts are vulnerable to disturbance and/or destruction (e.g., recreational caving or mine exploration, mine reclamation, and renewed mining in historic districts); not yet known to be afflicted by white-nose syndrome, but this fungal disease of bats now occurs throughout much of the eastern portion of the range of C. townsendii; currently not known to be threatened by mortality caused by wind turbines, but this might pose a threat in the future.
Range Extent Comments
Range includes western North America from southern British Columbia south to the Isthmus of Tehuantepec (Mexico), west to the Pacific coast, eastward to the Black Hills of South Dakota and Edwards Plateau of Texas, with isolated populations in the gypsum caves of northeastern Texas, Oklahoma, and Kansas, and in limestone regions of Arkansas, Missouri, Illinois, Indiana, Ohio, Kentucky, Virginia, and West Virginia. Elevational range extends from near sea level to at least 3,300 meters in some areas.

According to Handley (1959), the range of subspecies townsendii includes southwestern British Columbia, western Washington, western Oregon, and northwestern and west-central California; this subspecies intergrades with subspecies pallescens over a wide area in central and northern California and northward between the Cascades and the Rockies; subspecific allocation of specimens from much of this area is uncertain; some authors have applied the subspecific name intermedius to populations in the area of intergradation (Handley 1959). Handley characterized the range of subspecies pallescens as extending from southern British Columbia southward through the western United States to northwestern Mexico; west to central Washington, central Oregon, eastern and southern California, Sonora, and islands in the Gulf of California; east to South Dakota, Kansas, western Oklahoma, and Texas. As defined by Handley, subspecies australis ranges from the mountains of central and northern Mexico to western Texas.

However, a range-wide study of molecular phylogeny by Piaggio and Perkins (2005) found that geographic patterns of genetic (DNA) variation do not conform with the ranges of subspecies townsendii and pallescens as defined by Handley based on morphological characteristics. Piaggio and Perkins (2005) concluded that subspecies townsendii is broadly distributed from the Pacific coast of the United States and British Columbia (including Vancouver Island) southward to coastal regions of the Sonoran Desert in Mexico, eastward to the Colorado Plateau and the Black Hills, whereas subspecies pallescens is much more restricted and ranges from New Mexico to central and eastern Colorado, where it reaches its northern limits in northern Colorado or southern Wyoming. The molecular phylogenetic results conformed with the geographic scope of australis as defined by Handley.

Subspecies ingens: Range includes the Ozark Highlands and Boston Mountains ecoregions in northeastern Oklahoma, northwestern Arkansas, and (at least formerly) southwestern Missouri.

Subspecies virginianus: Range encompasses the Appalachian Mountains in Virginia, West Virginia, North Carolina, and eastern Kentucky. Presently these bats occur in decreased numbers throughout much of the historical range. The largest colonies are in several caves in Pendleton County, West Virginia; some caves serve as both hibernation and maternity sites, others are primarily maternity caves. Colonies occur also in Lee County and surrounding counties, Kentucky (the best known site being Stillhouse Cave); in Bath, Highland, Rockingham, Bland, and Tazewell counties, Virginia (Dalton 1987); and in Avery and Watauga counties, North Carolina (including Black Rock Cliffs Cave) See Matthews and Moseley (1990) and Handley (1991).

The molecular phylogeny of Piaggio and Perkins (2005) agreed with the morphologically defined taxonomic/geographic scope of subspecies ingens and virginianus as defined by Handley (1959).
Occurrences Comments
The number of distinct occurrences has not been determined using standardized criteria, but this species is represented by a very large number of known roost/observation sites and locations (as defined by IUCN).

These bats roost pendant-like on open surfaces of caves and abandoned mines (Western Bat Working Group 2005). Thus it is relatively unlikely to be missed in surveyed areas of caves and abandoned mines, even when present in low numbers (as is typical). However, recent studies indicate that use of roost sites is variable within seasons and among years, so multiple surveys may be required before use can be documented within an area (Western Bat Working Group 2005). Also, roosts can be very difficult to locate in some regions (particularly Canada and some desert areas) (Western Bat Working Group 2005).

Since 2008, the Subterranean Program of Bat Conservation International (BCI) has surveyed thousands of mines and caves across the United States, with a focus on the western and southwestern United States. During these surveys C. townsendii consistently was the most encountered bat. On BLM lands in the western United States, BCI surveyed 1,659 abandoned mine features and encountered a total of 211 C. townsendii in approximately 127 mines (K. Gillies, pers. comm., 2015).

In British Columbia, this species is represented by about 100 occurrences (not defined, but presumably based on roost sites) (Nagorsen 2013). In the 1990s, this species was represented in California by 37 known colonies (Pierson and Rainey 1998). In northern Utah, big-eared bats were found in day roosts in 196 caves and mines (none of 105 bridges) out of 676 mines and 39 caves surveyed; 13 were maternity roosts (Sherwin et al. 2000). In southwestern Colorado, Hayes et al. (2011) found C. corynorhinus in 38 of 133 mines surveyed. Townsend's big-eared bats were found in 9 of 25 abandoned mines surveyed in Durango, Mexico (López-González and Torres-Morales 2004).

Subspecies ingens: As of 2008, this subspecies was represented by 20 essential caves, including 10 maternity colonies, 8 hibernacula, and 2 used only in autumn; several caves are used both as a maternity site and hibernaculum (USFWS 2008). Several essential caves serve as alternate roosts for the same maternity colony (colony may move among the alternate caves during both the maternity season and between years) (USFWS 2008a). Same-year counts at maternity sites and hibernacula indicate that it is likely that major hibernacula have not yet been located (USFWS 2008a).

Subspecies virginianus: Currently known from 13 major (more than 200 bats) maternity colonies (USFWS 2008b).
Threat Impact Comments
The primary threat appears to be disturbance and/or destruction of roost sites resulting from: recreational caving or mine exploration; mine reclamation; renewed mining in historic districts; destruction/decay of buildings used as roosts, or reuse by people or deliberate exclusion of bats from such buildings (Western Bat Working Group 2005, Hayes and Wiles 2013)."In large portions of its western range, dependence upon abandoned mines puts this species at risk if mine reclamation and renewed mining projects do not mitigate for roost loss, or do not conduct adequate biological surveys prior to mine closure" (Western Bat Working Group 2005).

"Surveys conducted in Oregon and California indicate that current and historic roost sites have been negatively impacted by human visitation and renewed mining in recent years with most reported colonies exhibiting moderate to sizable reduction in numbers. Additional surveys in Utah indicate that several historic maternity sites have been abandoned, although it is not known if these colonies have relocated." Source: Western Bat Working Group 2005).

Nondestructive intrusion associated with research activities can negatively affect local colonies (Pearson et al. 1952), and capture and handling of bats in nursery roosts may cause the bats to vacate the site (Humphrey and Kunz 1976). Archeological excavations in Stanton’s Cave, Grand Canyon National Park, clearly caused roost abandonment. This cave was fenced for a few years and considerable activity in the front chamber caused high levels of disturbance in the form of in and out traffic, noise, and light (K. Gillies, pers. comm., 2015). Despite this sensitivity, careful nondestructive intrusion by researchers usually is not a significant problem; day-roosting individuals (López-González and Torres-Morales 2004) and females in maternity sites generally appear to be somewhat tolerant of disturbance (Sherwin et al. 2000; K. Gillies, pers. comm., 2015).

Both roosting and foraging areas may be negatively impacted by timber harvest practices and loss of riparian habitat (Western Bat Working Group 2005). See also Piaggio et al. (2009).

Activities associated with renewed oil exploration and extraction could negatively affect some populations of subspecies virginianus (see USFWS 2008b).

Since this species feeds largely on moths, pesticide spraying to control outbreaks of moth pests (e.g., spruce budworm, tussock moths, and Spongy Moth - Lymantria dispar) and other insects on forest and agricultural lands near roosts may affect overall moth abundance and might reduce food resources for this species (Hayes and Wiles 2013, Nagorsen 2013). However, population-level impacts of such pesticide use on this species are uncertain.

Lack of long-term protection is a conservation concern in some areas within the range of subspecies virginianus. Habitat loss and degradation from human activities are ongoing concerns, and all populations are vulnerable to significant losses from predation or vandalism (USFWS 2008b)

White-nose syndrome has not yet been documented as afflicting Townsend's big-eared bats, but this fungal disease of bats now occurs throughout much of the eastern portion of the range of C. townsendii, including caves used by C. townsendii (Stihler 2011a).

Mortality associated with wind turbines is a potential threat. Currently, turbines are not known to be a significant source of mortality (e.g., see USFWS 2008a, b), but they could become so as more turbines are installed throughout the range of the species (Miller et al. 2011).

Piaggio et al. (2009) found that population structuring estimated from mtDNA and microsatellites showed significant levels of differentiation among populations of C. t. virginianus located in different geographical regions. The authors stated that these levels of regional differentiation suggest a complete loss of connectivity among regional populations of C. t. virginianus among females and among males except between the northeastern and central West Virginia regions. Further, microsatellite data support loss of connectivity of these regional populations through evidence of population bottlenecks and inbreeding in some populations of C. t. virginianus (Piaggio et al. 2009).

Neither western subspecies (C. t. pallescens, C. t. townsendii) shows signs of a reduction in population genetic diversity (Piaggio et al. 2009).

Western subspecies townsendii and pallescens: The greatest threat is vandalism, and disturbance by humans; disturbance of a nursery colony or of a hibernating group may sometimes cause the bats to abandon the site and move to an alternate roost (Pearson 1952, Handley 1959). An additional threat is blockage of cave/mine entrances through collapse or human activities, including mine closures related to human safety issues (Perkins 1990; pers. comm., 1997; Wyoming State Wildlife Action Plan 2010). Habitat loss is also a threat.

Subspecies ingens: The historical decline probably was attributable mainly to human intrusion into caves, which depletes energy reserves of aroused bats and may lead to cave abandonment if disturbance is frequent (Pearson 1952, Handley 1959). Females exhibit limited dispersal, and individuals exhibit high natal philopatry, so extirpated colony site are unlikely to be naturally recolonized (Weyandt et al. 2005, USFWS 2008a). The occupied region is experiencing considerable human population growth and associated development; as a result, big-eared bat populations may be subject to increasing habitat loss and fragmentation, vandalism, and distubance (USFWS 2008a). Recreational use and associated human disturbance at maternity caves and hibernacula remain a major threat (USFWS 2008a). Likely there are as yet undiscovered major roosts, and threats to these are unknown (USFWS 2008a). Currently, disease and predation are not considered major threats (USFWS 2008a).

Subspecies virginianus: The growing popularity of spelunking is a tremendous threat to these bats. Very intolerant of disturbance in summer and winter. Former decline probably is attributable to human intrusion into caves, which depletes energy reserves of aroused bats and may lead to cave abandonment if disturbance is frequent. Forest defoliation by the introduced Spongy Moth (Lymantria dispar) could adversely affect native Lepidoptera and impact bat population (Sample and Whitmore 1993). In some areas, feral cats prey on the bats.

A relatively low degree of genetic diversity has been identified in populations of the endangered eastern subspecies, C. t. virginianus as inferred from both mtDNA and microsatellite DNA (Piaggio et al. 2009). "This reduced genetic diversity means that genetic drift may be driving diversity within these populations and the biodiversity and evolutionary potential of C. t. virginianus has been diminished" (Piaggio et al. 2009).
Ecology & Habitat

Description

Very large ears, 30-39 mm, joined across forehead; dorsal hairs slate or gray with pale cinnamon brown to blackish brown tips that contrast little with the base; ventral hairs slate, gray, or brownish, with brownish or buff tips; two large fleshy lumps on snout; hairs on toes do not project beyond toenails; total length 90-112 mm; forearm 39.2-47.6 mm; greatest length of skull 15.2-17.4 mm; 36 teeth; adult mass 5-13 g (Handley 1959, Hall 1981, Kunz and Martin 1982, Ingles 1965).

Diagnostic Characteristics

Differs from Idionycteris phyllotis in having a posteriorly elongated nostril, a conspicuous glandular mass on each side of the muzzle between the eye and nostril, lack of an accessory basal lobe of the ear, an unkeeled calcar, and breadth of braincase less than half the greatest length of the skull (Handley 1959, Hall 1981). Differs from Corynorhinus rafinesquii in lacking sharp contrast between the bases and tips of the ventral hairs (tips white or whitish in rafinesquii) and by having a strong undepressed rostrum. Differs from Corynorhinus mexicanus in having a longer skull (usually more than 15.5 mm or 15.7 mm vs. usually less than 15.5 mm or 15.7 mm, for males and females, respectively), longer tragus (usually more than 13 mm vs. usually less than 13 mm), longer maxillary toothrow (usually more than 4.9 mm vs. usually less than 4.9 mm), and more interfemoral croos ribs (usually more than 9 vs. usually fewer than 9) (Handley 1959, Hall 1981, Jones 1977). See Jones (1977) for further dental and cranial distinctions.

Habitat

Throughout much of the known range, these bats commonly occur in mesic habitats characterized by coniferous and deciduous forests (Kunz and Martin 1982), but they occupy a broad range of habitats (e.g., see Handley 1959). Generally they are uncommon in prairies and extreme desert, although they occur in the lower elevations of the arid plateau and desert ranges of northcentral Mexico and the arid valleys south of the transverse volcanic belt. They are known in Mexico mostly from relatively arid regions (e.g., grassy hills with nearby pine-oak woodland) but also from more humid localities with oak, pine, juniper, cypress, madrone, and manzanita (Handley 1959). Ozark and Appalachian populations inhabit caves mostly in oak-hickory forest (Handley 1959).

On the West Coast, Townsend's big-eared bats are found regularly in forested regions and buildings, and in areas with a mosaic of woodland, grassland, and/or shrubland. In California and Washington, they are known from limestone caves, lava tubes, and human-made structures in coastal lowlands, cultivated valleys, and nearby hills covered with mixed vegetation (Handley 1959). Fellers and Pierson (2002) studied a population in Point Reyes National Seashore, California, that foraged along the edges of redwood and Douglas-fir forests and woodlands, primarily along the edges of riparian vegetation. Individuals avoided grassland wherever possible, even while commuting to foraging grounds. Individuals hunted primarily around the perimeter of trees, usually 10-30 meters off the ground, between mid-canopy and near the top of the canopy (Fellers and Pierson 2002).

Townsend's big-eared bats are recorded from pine-fir-hemlock-broadleaf deciduous forest in western Oregon, and from the edge of spruce-fir forest in Colorado (see Handley 1959). In Utah, day roosts were associated with sagebrush steppe, juniper woodlands and mountain brush vegetation at lower available elevations (1,350-2,440 meters; Sherwin et al. 2000). In Texas, habitat ranges from desert scrub to pinyon-juniper woodland, consistently in areas with canyons or cliffs (Schmidly 1991). In New Mexico, most have been captured in evergreen forests during warm months, least commonly captured in xeric shrublands (see Kunz and Martin 1982). In Arizona, habitats include desertscrub, in shelters in desert mountains (where infrequent), oak woodland, pinyon-juniper, and conifer forests (Hoffmeister 1986). In Kansas and Oklahoma, these bats are apparently restricted to riparian communities and nearby gypsum caves in the mid-grass prairie region. In Oklahoma, foraging adult females used edge habitats of intermittent streams and mountain slopes more than expected based on relative availability of habitats (Clark et al. 1993). During the prehibernation period in Oklahoma, Wethington et al. (1996) found that females used habitats in proportion to their availability, but males used forested habitat more than expected in September, likely reflecting prey availability. Females in West Virginia used a wide range of foraging habitats, including deciduous forest, mixed deciduous/ coniferous forest, old fields, hayfields, and corn fields (Stihler 2011).

Maternity and hibernation colonies typically are in caves and mine tunnels. These bats prefer relatively cold places for hibernation, often near entrances and in well-ventilated areas. In California, most limestone caves are too warm for successful hibernation; solitary males and small groups of females are known to hibernate in buildings in the central part of the state. These bats do not use crevices or cracks; they hang from the ceiling, generally near the zone of total darkness (Schmidly 1991). In some areas, basal hollows in large trees may as roost sites, especially in areas with few caverns (Fellers and Pierson 2002, Mazurek 2004). In western Colorado, C. townsendii hibernated usually in abandoned mines with more than one opening and with portal temperatures near 0 C, though they sometimes used mines with a single opening and much warmer portal temperatures (Hayes et al. 2011).

Females gather in small nursery colonies in the warm parts of caves or mines, and not uncommonly in buildings (western North America). Individuals generally return to the same maternity roost and hibernaculum in successive years, though individuals may move among roosts during both the maternity season and in winter (Clark et al. 2002). Night roosts include caves, buildings, and tree cavities.

Maternity roosts in British Columbia exist in buildings and caves (Reid et al. 2010). In coastal California, all six known maternity colonies were in old buildings (5) or in a cave-like feature of a bridge (1) (Fellers and Pierson 2002). In northwestern California, a maternity colony was in a basal hollow of a live coast redwood; this population may have moved among multiple tree hollows in the area (Mazurek 2004). In Oregon, both sexes apparently use a series of interim roost sites between emergence from hibernation and the time females enter maternity colonies, with little individual fidelity to these sites (Dobkin et al. 1995). In Utah, caves were preferred as day roosts in summer (85% of surveyed caves used), as well as abandoned mines (21% surveyed were used); no bridges were used (Sherwin et al. 2000). In Oklahoma-Arkansas-Missouri, big-eared bats are obligate cave users year round; they are known to utilize and roost in limestone and sandstone talus caves (USFWS 2008).

Ecology

Crude population density in Oklahoma was estimated at one bat per 46.6 sq. km (see Kunz and Martin 1982), about 3-4 times greater than that reported for populations in California (Pearson et al. 1952).

These bats hibernate singly, or in clusters in some areas (Caire et al. 1989, Schmidly 1991). They tend not to associate with other species of bats in daytime or in hibernation roosts, though scattered individuals of other normally colonial species occasionally may be present (Handley 1959).

Pre-weaning post-natal mortality generally is low. Adult survivorship is relatively high (about 70-80% in females in California). Estimates of annual survival of wintering Townsend's big-eared bats in 3 locations in Washington ranged from 54 to 76.0 percent and varied by location, time or trends, and sex (Ellison 2010)..

Predation has been suggested as the primary limiting factor in Kansas and Oklahoma (see Handley 1959).

Pearson et al. (1952) believed that a population increase might be dependent on the establishment of new nursery colonies (colonies remained static in size year after year); how new nursery colonies become established is not known (Handley 1959).

Reproduction

Mating begins in autumn, continues into winter. Ovulation and fertilization are delayed until late winter/early spring. Gestation lasts 2-3.5 months. A litter of one is born in late spring/early summer (beginning mainly in late May in California, the second week of July in Washington, and June in southwestern Texas). Throughout the U.S. range, the earliest births occur in mid-April, the latest in late July (see Handley 1959). Young can fly at 2.5-4 weeks and are weaned by 6-8 weeks. Females are sexually mature their first summer. Males are not sexually active until their second year (California). Nearly all adult females breed every year. Maximum longevity exceeds 21 years (Perkins 1994).

Females commonly form nursery colonies generally of up to about 200 (west) or 1,000 (east) individuals, but solitary pregnant females are frequently encountered (Handley 1959); males roost separately (apparently solitarily) during this time.
Terrestrial Habitats
Forest/WoodlandForest - HardwoodForest - ConiferForest - MixedForest EdgeWoodland - HardwoodWoodland - ConiferWoodland - MixedShrubland/chaparralSavannaGrassland/herbaceousOld fieldDesertCliff
Palustrine Habitats
Riparian
Other Nations (2)
CanadaN3
ProvinceRankNative
British ColumbiaS3Yes
United StatesN3
ProvinceRankNative
KansasS2Yes
West VirginiaS2Yes
OregonS2Yes
MontanaS3Yes
NevadaS2Yes
UtahS3Yes
MissouriSXYes
IdahoS3Yes
CaliforniaS2Yes
WyomingS2B,S1NYes
ColoradoS2Yes
NebraskaS1Yes
TexasS3Yes
WashingtonS3Yes
New MexicoS3Yes
North CarolinaS1Yes
North DakotaSNRYes
ArizonaS3Yes
VirginiaS1Yes
Navajo NationS2Yes
South DakotaS2Yes
ArkansasS1Yes
KentuckySNRYes
OklahomaS3Yes
TennesseeSNRYes
Threat Assessments
ThreatScopeSeverityTiming
1 - Residential & commercial developmentNegligible (<1%)High (continuing)
2 - Agriculture & aquacultureNegligible (<1%)High (continuing)
2.1 - Annual & perennial non-timber cropsNegligible (<1%)High (continuing)
3 - Energy production & miningRestricted - smallModerate - slightHigh (continuing)
3.2 - Mining & quarryingSmall (1-10%)Moderate - slightHigh (continuing)
3.3 - Renewable energySmall (1-10%)Slight or 1-10% pop. declineHigh (continuing)
4 - Transportation & service corridorsNegligible (<1%)High (continuing)
5 - Biological resource useSmall (1-10%)Moderate - slightHigh (continuing)
5.1 - Hunting & collecting terrestrial animalsSmall (1-10%)Moderate - slightHigh (continuing)
5.3 - Logging & wood harvestingSmall (1-10%)Moderate - slightHigh (continuing)
6 - Human intrusions & disturbanceRestricted - smallSlight or 1-10% pop. declineHigh (continuing)
6.1 - Recreational activitiesHigh (continuing)
6.3 - Work & other activitiesHigh (continuing)
7 - Natural system modificationsNegligible (<1%)High (continuing)
7.1 - Fire & fire suppressionHigh (continuing)
7.2 - Dams & water management/useHigh (continuing)
8 - Invasive & other problematic species, genes & diseasesLarge - restrictedNegligible or <1% pop. declineHigh (continuing)
9 - PollutionSmall (1-10%)Slight or 1-10% pop. declineHigh (continuing)
10 - Geological eventsNegligible (<1%)
11 - Climate change & severe weatherPervasive (71-100%)UnknownHigh (continuing)

Roadless Areas (41)
Arizona (2)
AreaForestAcres
HellsgateTonto National Forest6,171
WhetstoneCoronado National Forest20,728
California (27)
AreaForestAcres
AgnewSequoia National Forest9,561
Big RocksLos Padres National Forest11,866
Bucks LakePlumas National Forest680
Callahan FlowModoc National Forest6,618
ChanchelullaShasta-Trinity National Forest3,915
ChicoSequoia National Forest39,836
Chips CreekPlumas National Forest12,940
Coyote NorthInyo National Forest11,932
Dobie FlatModoc National Forest15,079
Eagle PeakCleveland National Forest6,481
Ferguson RidgeSierra National Forest6,104
Garcia MountainLos Padres National Forest7,850
Hoover - NorthHumboldt-Toiyabe National Forest1,574
Iceberg - Mill CreekHumboldt-Toiyabe National Forest26,988
KangarooKlamath National Forest40,617
LavasModoc National Forest25,864
Lion RidgeSequoia National Forest5,265
Middle ForkPlumas National Forest29,278
MonarchSierra National Forest697
Mt. HoffmanModoc National Forest9,780
No NameCleveland National Forest4,897
RinconSequoia National Forest54,610
Salt GulchShasta-Trinity National Forest6,511
South ForkShasta-Trinity National Forest16,786
SugarloafSan Bernardino National Forest8,206
Trumbull PeakStanislaus National Forest6,164
West GirardShasta-Trinity National Forest37,516
Montana (2)
AreaForestAcres
Middle Mtn. / Tobacco RootsBeaverhead-Deerlodge National Forest96,487
Scotchman Peaks (MT)Kootenai National Forest53,909
Nevada (1)
AreaForestAcres
Boundary Peak (NV)Inyo National Forest21,851
New Mexico (3)
AreaForestAcres
Capitan MountainsLincoln National Forest14,069
Frisco BoxGila National Forest38,979
Peloncillo (NM)Coronado National Forest43,339
Oregon (1)
AreaForestAcres
HomesteadWallowa-Whitman National Forest5,817
Utah (1)
AreaForestAcres
Temple PeakWasatch-Cache National Forest24,081
Washington (3)
AreaForestAcres
Laughing WaterGifford Pinchot National Forest1,167
LightningOlympic National Forest7,179
ManastashWenatchee National Forest11,155
Wyoming (1)
AreaForestAcres
Middle ForkShoshone National Forest51,772
References (97)
  1. American Society of Mammalogists (ASM). 2025. Mammal Diversity Database (Version 1.13) [Data set]. Zenodo. https://doi.org/10.5281/zenodo.10595931. Online. Available: https://www.mammaldiversity.org/
  2. Ammerman, L. K., C. L. Hice, and D. J. Schmidly. 2012. Bats of Texas. Texas A &amp; M University Press, College Station, Texas. xvi + 305 pp.
  3. Arita, H. T. 1993. Conservation biology of the cave bats in Mexico. Journal of Mammalogy 74:693-702.
  4. Avila-Flores, R. and R. A. Medellín. 2004. Ecological, taxonomic, and physiological correlates of cave use by Mexican bats. Journal of Mammalogy 85:675-687.
  5. Bagley, F. M. 1984. Recovery plan for the Ozark big-eared bat and the Virginia big-eared bat. U.S. Fish and Wildlife Service. 119 pp.
  6. Bagley, R. and J. Jacobs. 1985. Census techniques for endangered big-eared bats proving successful. Endangered Species Tech. Bull. 10(3):5-7.
  7. Baker, R. J., L. C. Bradley, R. D. Bradley, J. W. Dragoo, M. D. Engstrom, R. S. Hoffman, C. A. Jones, F. Reid, D. W. Rice, and C. Jones. 2003a. Revised checklist of North American mammals north of Mexico, 2003. Museum of Texas Tech University Occasional Papers 229:1-23.
  8. Barbour, R. W., and W. H. Davis. 1969. Bats of America. The University of Kentucky Press, Lexington, Kentucky. 286 pp.
  9. Bayless, M. L., M. K. Clark, R. C. Stark, B. S. Douglas, and S. M. Ginger. 2011. Distribution and status of eastern big-eared bats (<i>Corynorhinus </i>spp.). Pages 13-25 in Loeb, S.C., M. J. Lacki, and D. A. Miller, editors. Conservation and managementof big-eared bats: a symposium. USDA Forest Service, Southern Research Station, Ashville.
  10. Betts, B. J. 2010. Activity budgets of Townsend's big-eared bats at a maternity colony in northeastern Oregon. Northwestern Naturalist 91:13-22.
  11. Bogdanowicz, W., S. Kasper, and R. D. Owen. 1998. Phylogeny of plecotine bats: reevaluation of morphological and chromosomal data. Journal of Mammalogy 79:78-90.
  12. Bradley, R.D., L.K. Ammerman, R.J. Baker, L.C. Bradley, J.A. Cook. R.C. Dowler, C. Jones, D.J. Schmidly, F.B. Stangl Jr., R.A. Van den Bussche and B. Würsig. 2014. Revised checklist of North American mammals north of Mexico, 2014. Museum of Texas Tech University Occasional Papers 327:1-28. Available at: http://www.nsrl.ttu.edu/publications/opapers/ops/OP327.pdf
  13. Burford, L. S., and M. J. Lacki. 1995. Habitat use by <i>Corynorhinus townsendii virginianus</i> in the Daniel Boone National Forest. American Midland Naturalist 134:340-345.
  14. Caire, W., J. D. Tyler, B. P. Glass, and M. A. Mares. 1989. Mammals of Oklahoma. University of Oklahoma Press, Norman. Oklahoma. 567 pp.
  15. Choate, J. R., and J. M. Anderson. 1997. Bats of Jewel Cave National Monument, South Dakota. Prairie Naturalist 29:39-47.
  16. Clark, B. K., B. S. Clark, and D. M. Leslie, Jr. 1997a. Seasonal variation in use of caves by the endangered Ozark big-eared bat (<i>Corynorhinus townsendii ingens</i>) in Oklahoma. American Midland Naturalist 137:388-392.
  17. Clark, B. S., B. K. Clark, and D. M. Leslie, Jr. 2002. Seasonal variation in activity patterns of the endangered Ozark big-eared bat (<i>Corynorhinus townsendii ingens</i>). Journal of Mammalogy 83:590-598.
  18. Clark, B. S., D. M. Leslie, Jr., and T. S. Carter. 1993. Foraging activity of adult female Ozark big-eared bats (PLECOTUS TOWNSENDII INGENS) in summer. J. Mammalogy 74:422-427.
  19. Clark, S. C., W. L. Puckette, B. K. Clark, and D. M. Leslie, Jr. 1997b. Status of the Ozark big-eared bat (<i>Corynorhinus townsendii ingens</i>) in Oklahoma, 1957 to 1995. Southwestern Naturalist 42:20-24.
  20. Clawson, R. L., R. K. LaVal, M. L. LaVal, and W. Claire. 1980. Clustering behavior of hibernating <i>Myotis sodalis</i> in Missouri, USA. Journal of Mammalogy 61:245-53.
  21. Dalton, V.M. 1987. Distribution, abundance, and status of bats hibernating in caves in Virginia. Virginia Journal of Science 38(4): 369-379.
  22. Dobkin, D. S., R. D. Gettinger, and M. G. Gerdes. 1995. Springtime movements, roost use, and foraging activity of Townsend's big-eared bat (PLECOTUS TOWNSENDII) in central Oregon. Great Basin Naturalist 55:315-321.
  23. Dodd, L. 2006. Diet and prey abundance of the Ozark big-eared bat (<i>Corynorhinus townsendii ingens</i>) in Arkansas. Master's thesis. University of Kentucky, Lexington, Kentucky. 252 pp.
  24. Elliott, W. R., K. B. Lister, and M. A. Shiver. 1999. A survey for Ozark big-eared bats <i>Corynorhinus townsendii ingens</i> and a cave crayfish <i>Cambarus aculabrum</i> in southern Missouri. Missouri Department of Conservation. Unpublished report. 29 pp.
  25. Ellison, L. E. 2010. A retrospective survival analysis of Townsend's big-eared bat (<i>Corynorhinus townsendii</i>) from Washington state. Northwestern Naturalist 91:172-182.
  26. Fellers, G. M., and E. D. Pierson. 2002. Habitat use and foraging behavior of Townsend's Big-eared Bat (<i>Corynorhinus townsendii</i>) in coastal California. Journal of Mammalogy 83:167-177.
  27. Frost, D. R., and R. M. Timm. 1992. Phylogeny of plecotine bats (Chiroptera: "Vespertilionidae"): proposal of a logically consistent taxonomy. Am. Mus. Novitates 3034:1-16.
  28. Garner, J. D., and J. E. Gardner. 1992. Determination of summer distribution and habitat utilization of the Indiana bat (<i>Myotis sodalis</i>) in Illinois. Illinois Department of Conservation. Final Report, Project E-3. Springfield, IL, 23 pp.
  29. Graening, G.O., M. J. Harvey, W. L. Puckette, et al. 2011. Conservation status of the endangered Ozark big-eared bat (<i>Corynorhinus townsendii ingens</i>)--a 34-year assessment. Publications of the Oklahoma Biological Survey. 11:1-16.
  30. Griffin, D. R. 1940b. Migrations of New England bats. Bulletin of the Museum of Comparative Zoology 86:217-246.
  31. Gruver, J. C., and D. A. Keinath. 2006. Townsend's big-eared bat (<i>Corynorhinus townsendii</i>): A technical conservation assessment. Report to the U.S. Department of Agriculture Forest Service, Rocky Mountain Region. Available at http://www.fs.fed.us/r2/projects/scp/assessments/townsendsbigearedbat.pdf.
  32. Hall, E. R. 1981a. The Mammals of North America, second edition. Vols. I &amp; II. John Wiley &amp; Sons, New York, New York. 1181 pp.
  33. Hamilton, W. J., Jr., and J. O. Whitaker, Jr. 1979. Mammals of the eastern United States. Cornell Univ. Press, Ithaca, New York. 346 pp.
  34. Handley, C. O., Jr. 1959. A revision of American bats of the genera Euderma and Plecotus. Proceedings U.S. National Museum 110:95-246.
  35. Handley, C. O., Jr. 1991. Mammals. Pages 539-616 in K. Terwilliger, coordinator. Virginia's endangered species: proceedings of a symposium. McDonald and Woodward Publishing Company, Blacksburg, Virginia.
  36. Hardy, R. 1941. Some notes on Utah bats. J. Mammal. 22: 289-295.
  37. Hasenyager, R. N. 1980. Bats of Utah. Utah State Division of Wildlife Resources, Salt Lake City. Publ. No. 80-15, 109 pp.
  38. Hayes, G., and G. J. Wiles. 2013. Washington bat conservation plan. Washington Department of Fish and Wildlife, Olympia, Washington. viii + 138 pp.<br>
  39. Hayes, M. A., R. A. Schorr, and K. W. Navo. Hibernacula selection by Townsend's big-eared bat in southwestern Colorado. Journal of Wildlife Management 75:137-143.
  40. Hensley, S. 2003. The treasures of the Ozark Plateau. Endangered Species Bulletin 28(1):32-33.
  41. Humphrey, S.R. and T.H. Kunz. 1976. Ecology of a Pleistocene relic, the western big-eared bat (<i>Plecotus townsendii</i>), in the southern Great Plains. Journal of Mammalogy 57(3):470-494.
  42. Ingles, L. G. 1965. Mammals of the Pacific States. Stanford University Press, Stanford, California.
  43. Jones, C. 1977. Plecotus rafinesquii. American Soc. Mamm., Mammalian Species No. 69. 4 pp.
  44. Kiser, J. D. and C. L. Elliott. 1996. Foraging habitat, food habits, and roost tree characteristics of the Indiana bat (<i>Myotis sodalis</i>) during autumn in Johnson County, Kentucky. Final report, Kentucky Department of Fish and Wildlife Resources, Frankfort, Kentucky. 65pp.
  45. Kunz, T.H. and R.A. Martin. 1982. <i>Plecotus townsendii</i>. American Society Mamm., Mammalian Species No. 175. 6 pp.
  46. Lacki, M. J., and L. E. Dodd. 2011. Diet and foraging behavior of <i>Corynorhinus </i>in eastern North America. Pages 39-52 in Loeb et al., editors. Conservation and management of eastern big-eared bats. USDA Forest Service, Southern Research Station, General Technical Report SRS-145.
  47. Leslie, D. M., and B. S. Clark. 2002. Feeding habits of the endangered Ozark big-eared bat (<i>Corynorhinus townsendii ingens</i>) relative to prey abundance. Acta Chiropterologia 4(2):173-182.
  48. Lewis, L. 1994a. Assessment of bat inventory and monitoring data in the Shoshone District BLM. Bureau of Land Management, Shoshone District, Shoshone, Idaho.
  49. Linzey, D. W. 1998. The mammals of Virginia. McDonald and Woodward, Blacksburg, VA. 459 pp.
  50. Loeb, S. C., M. J. Lacki, and D. A. Miller, editors. 2011. Conservation and management of eastern big-eared bats. USDA Forest Service, Southern Research Station, General Technical Report SRS-145.
  51. López-González, C., and L. Torres-Morales. 2004. Use of abandoned mines by long-eared bats, genus <i>Corynorhinus </i>(Chiroptera: Vespertilionidae) in Durango, Mexico. Journal of Mammalogy 85:989-994.
  52. Mammalian Species, nos. 1-604. Published by the American Society of Mammalogists.
  53. Matthews, J.R. and C.J. Moseley (eds.). 1990. The Official World Wildlife Fund Guide to Endangered Species of North America. Volume 1. Plants, Mammals. xxiii + pp 1-560 + 33 pp. appendix + 6 pp. glossary + 16 pp. index. Volume 2. Birds, Reptiles, Amphibians, Fishes, Mussels, Crustaceans, Snails, Insects, and Arachnids. xiii + pp. 561-1180. Beacham Publications, Inc., Washington, D.C.
  54. Mazurek, M. J. 2004. A maternity-roost of Townsend's big-eared bats (<i>Corynorhinus townsendii</i>) in coast redwood basal hollows in northwestern California. Northwestern Naturalist 85:60-62.
  55. Menzel, M. A., R. Odom, S. Owen, W. M. Ford, B. R. Chapman, K. V. Miller, J. Edwards, and P. Wood. 1999b. Investigation of foraging habitat use by bats with a focus on Northern Long-eared Myotis (<i>Myotis septentrionalis</i>): a comparison of methods. IN M. K. Clark, editor. Abstracts from the 1999 Colloquium on the conservation of mammals in the Southeastern United States. Available at: http://www.batworkinggroups.org/sbdnnews.htm. Accessed 2001-06-12.
  56. Miller, D. A., C. W. Stihler, D. B. Sasse, R. Reynolds, P. Van Deusen, and S. B. Castleberry. 2011. Conservation and management of eastern big-eared bats (<i>Corynorhinus </i>spp.). Pages 53-61 in Loeb et al., editors. Conservation and management of eastern big-eared bats. USDA Forest Service, Southern Research Station, General Technical Report SRS-145.
  57. Miller, K.E.G. 2007. Connectivity, relatedness, and winter roost ecology of Townsend’s big-eared bat (<i>Corynorhinus townsendii</i>). Masters Thesis, Idaho State University, vii + 74 pp.
  58. Miner, K. L., and D. C. Stokes. 2005. Bats in the South Coast Ecoregion: status, conservation issues, and research needs. Pages 211-227 in USDA Forest Service General Technical Report PSW-GTR-195.
  59. Morgan, C. N., L. K. Ammerman, K. D. Demere, J. B. Doty, Y. J. Nakazawa, and M. R. Mauldin. 2019. Field identification key and guide for bats of the United States of America. Occasional Papers, Museum of Texas Tech University, Number 360. Texas Tech University Natural Science Research Laboratory. 29 pp.
  60. Nagorsen, D. 2013, British Columbia Conservation Data Centre: conservation status report. <i>Corynorhinus townsendii</i>.
  61. Oliver, G. V. 2000. The bats of Utah: a literature review. Publication No. 00-14, Utah Division of Wildlife Resources, Salt Lake City.
  62. Pearson, O.P., M.R. Koford and A.K. Pearson. 1952. Reproduction of the lump-nosed bat (<i>Corynorhinus rafinesquii</i>) in California. J. Mamm. 33:273-320.
  63. Perkins, J.M. 1990. Results of population monitoring for the Category 2 species <i>Plecotus townsendii</i> in Oregon and Washington - 1989-1990. Contract #90-9-03.
  64. Perkins J. M. 1994. Longevity records for two vespertilionids. Bat Research News 35:79-80.
  65. Piaggio, A. J., and S. L. Perkins. 2005. Molecular phylogeny of North American long-eared bats (Vespertilionidae: <i>Corynorhinus</i>); inter- and intraspecific relationships inferred from mitochondrial and nuclear DNA sequences. Molecular Phylogenetics and Evolution 37:762-775.
  66. Piaggio, A. J., K. W. Navo, and C. W. Stihler. 2009. Intraspecific comparison of population structure, genetic diversity, and dispersal among three subspecies of Townsend's big-eared bats, <i>Corynorhinus townsendii townsendii</i>, <i>C. t. pallescens</i>, and the endangered <i>C. t. virginianus</i>. Conservation Genetics 10:143-159.
  67. Pierson, E. D., and W. E. Rainey. 1998b. Distribution, status, and management of Townsend's big-eared bat (<i>Corynorhinus townsendii</i>) in California. California Department of Fish and Game, Bird and Mammal Conservation Program Report 96-7, Sacramento, CA. Report submitted in 1994, updated and finalized in 1998.
  68. Pierson, E.D., M.C. Wackenhut, J.S. Altenback, P. Bradley, P. Call, D.L. Genter, C.E. Harris, B.L. Keller, B. Lengus, L. Lewis, B. Luce, K.W. Navo, J.M. Perkins, S. Smith, and L. Welch. 1999. Species conservation assessment and strategy for Townsend's big-eared bat (<i>Corynorhinus townsendii townsendii </i>and <i>Corynorhinus townsendii pallescens</i>). Idaho Conservation Effort, Idaho Department of Fish and Game, Boise, Idaho.
  69. Ports, M. A., and P. V. Bradley. 1996. Habitat affinities of bats from northeastern Nevada. Great Basin Naturalist 56:48-53.
  70. Prendergast, J. A., W. E. Jensen, and S. D. Roth. 2010. Trends in abundance of hibernating bats in a karst region of the southern Great Plains. Southwestern Naturalist 55:331-339.
  71. Reid, A., T. Hill, R. Clarke, J. Gwilliam and J. Krebs. 2010. Roosting ecology of female Townsend's Big-Earded Bats (<i>Corynorhinus townsendii</i>) in south-eastern British Columbia: Implications for Conservation Management. Northwestern Naturalist 91(2): 215-218.
  72. Schmidly, D. J. 1991. The bats of Texas. Texas A & M University Press, College Station, Texas. 188 pp.
  73. Schwartz, C. W., and E. R. Schwartz. 1981. The wild mammals of Missouri. University of Missouri Press, Columbia. 356 pp.
  74. Sherwin, R. E., D. Stricklan, and D. S. Rogers. 2000. Roosting affinities of Townsend's Big-eared Bat (<i>Corynorhinus townsendii</i>) in northern Utah. Journal of Mammalogy 81:939-947.
  75. Simmons, N. B. and A. L. Cirranello. 2025. Bat Species of the World: A taxonomic and geographic database. Version 1.7. Online. Available: https://batnames.org/
  76. Smith, S. J., D. M. Leslie, Jr., M. J. Hamilton, J. B. Lack, and R. A. Van Den Bussche. 2008. Subspecific affinities and conservation genetics of western big-eared bats (<i>Corynorhinus townsendii pallescens</i>) at the edge of their distributional range. Journal of Mammalogy 89:799-814.
  77. Stihler, C. W. 2011a. Status of the Virginia big-eared bat (<i>Corynorhinus townsendii virginianus</i>) in West Virginia: twenty-seven eyars of monitoring cave roosts. Pages 75-84 in Loeb et al., editors. Conservation and management of eastern big-eared bats. USDA Forest Service, Southern Research Station, General Technical Report SRS-145.
  78. Stihler, C. W. 2011b. Radiotelemetry studies of female Virginia big-eared bats (<i>Corynorhinus townsendii virginianus</i>) in Pendleton County, West Virginia. Pages 139-146 in Loeb et al., editors. Conservation and management of eastern big-eared bats. USDA Forest Service, Southern Research Station, General Technical Report SRS-145.
  79. Thomas, D. W. and S. D. West. 1989. Sampling methods for bats. USDA Forest Service, Pacific Northwest Research Station Gen. Technical Report PNW-GTR-243. 20 pp.
  80. Tumlison, R., and M. E. Douglas. 1992. Parsimony analysis and the phylogeny of the plecotine bats (Chiroptera: Vespertilionidae). Journal of Mammalogy 73(2):276-285.
  81. Tuttle, M. D. 1976a. Population ecology of the gray bat (<i>Myotis grisescens</i>): philopatry, timing, and patterns of movement, weight loss during migration, and seasonal adaptive strategies. University of Kansas Museum of Natural History Occasional Papers (54):1-38.
  82. U.S. Fish and Widllife Service (USFWS). 2009. Virginia big-eared bat (<i>Corynorhinus townsendii virginianus</i>) plan for controlled holding, propagation, and reintroduction. USFWS, West Virginia Field Office, Elkins, West Virginia.
  83. U.S. Fish and Wildlife Service (USFWS). 1999. Agency draft Indiana Bat (<i>Myotis sodalis</i>) revised recovery plan. U.S. Fish and Wildlife Service, Fort Snelling, Minnesota. 53 pp.
  84. U.S. Fish and Wildlife Service (USFWS). 2008a. Ozark big-eared bat (<i>Corynorhinus townsendii ingens</i>) 5-year review: summary and evaluation. USFWS, Oklahoma Ecological Services Field Office, Tulsa, OK.
  85. U.S. Fish and Wildlife Service (USFWS). 2008b. Virginia big-eared bat (<i>Corynorhinus townsendii virginianus</i>) 5-year review: summary and evaluation. Report prepared by West Virginia Field Office. 21 pp.
  86. U.S. Fish and Wildlife Service (USFWS). 2009. Virginia big-eared bat (<i>Corynorhinus townsendii virginianus</i>) plan for controlled holding, propagation, and reintroduction. USFWS, West Virginia Field Office, Elkins, WV.
  87. van Zyll de Jong, C.G. 1985. Handbook of Canadian Mammals. Vol. II, Bats. National Museum of Natural Sciences, National Museums of Canada, Ottawa, Canada. 212 pp.
  88. Wackenhut, M. C. 1990. Bat species overwintering in lava-tube caves in Lincoln, Gooding, Blaine, Bingham, and Butte counties, Idaho, with special reference to annual return of banded <i>Plecotus townsendii.</i> Unpublished thesis, Idaho State University, Pocate.
  89. Wainwright, J. M., and N. D. Reynolds. 2013. Cave hibernaculum surveys of a Townsend's big-eared bat (<i>Corynorhinus townsendii</i>) colony at Mount St. Helens, Washington. Northwestern Naturalist 94:240-244.
  90. Weller, T. J., S. C. Thomas. And J. A. Baldwin. 2014. Use of long-term opportunistic surveys to estimate trends in abundance of hibernating Townsend’s big-eared bats. Journal of Fish and Wildlife Management 5:59-69.
  91. Western Bat Working Group. 2005a. <i>Corynorhinus townsendii</i> Townsend's big-eared bat species account. Updated 2005.
  92. Wethington, T. A., D. M. Leslie, Jr., M. S. Gregory, and M. K. Wethington. 1996. Prehibernation habitat use and foraging activity by endangered Ozark big-eared bats (<i>Plecotus townsendii ingens</i>). American Midland Naturalist 135:128-230.
  93. Weyandt, S. E., R. A. Van Den Bussche, M. J. Hamilton, and D. M. Leslie, Jr. 2005. Unraveling the effects of sex and dispersal: Ozark big-eared bat (<i>Corynorhinus townsendii ingens</i>) conservation genetics. Journal of Mammalogy 86:1136-1143.
  94. White, D. H., and J. T. Seginak. 1987. Cave gate designs for use in protecting endangered bats. Wildlife Society Bull. 15:445-449.
  95. Wilson, D. E., and D. M. Reeder (editors). 1993. Mammal species of the world: a taxonomic and geographic reference. Second edition. Smithsonian Institution Press, Washington, DC. xviii + 1206 pp. Available online at: http://www.nmnh.si.edu/msw/.
  96. Wilson, D. E., and D. M. Reeder (editors). 2005. Mammal species of the world: a taxonomic and geographic reference. Third edition. The Johns Hopkins University Press, Baltimore. Two volumes. 2,142 pp. [As modified by ASM the Mammal Diversity Database (MDD) at https://www.mammaldiversity.org/index.html]
  97. Wilson, D. E., and S. Ruff. 1999. The Smithsonian book of North American mammals. Smithsonian Institution Press, Washington, D.C. 750 pp.