Myotis leibii are one of the rarest bats in North America. They occur from Ontario and Quebec in Canada southwestward through the Appalachian region to Arkansas and Eastern Oklahoma in the United States. They have been recorded in New England, Georgia, Alabama, Pennsylvania, Virginia, North Carolina, Kentucky, and Oklahoma. Fossil records have been found in Big Bone Cave in Tennessee and in Cumberland Cave in Maryland. The range of M. leibii is allopatric with M. californius, M. ciliolabrum and all other small footed Myotis bats with a keeled calcar in North America. (Best & Jennings 1997; Corgan 1975; Davis et al. 1965; Martin 1972; Wilson & Ruff 1999).

Biogeographic Regions: nearctic (Native )

occurs (regularly, as a native taxon) in multiple nations

Canada

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

Type of Residency: Year-round

United States

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

Type of Residency: Year-round

Global Range: (200,000-2,500,000 square km (about 80,000-1,000,000 square miles)) The range extends from New England, southeastern Ontario, and southwestern Quebec south and west to southeastern Oklahoma, Arkansas, northern Alabama, northern Georgia, and northwestern South Carolina (Menzel et al. 2003); see map in Best and Jennings (1997). Within this range, the distribution is very spotty, and the bulk of the occurrences and largest populations are in New York, Pennsylvania, West Virginia, and western Virginia. Hall's (1981) map should be taken as potential range; there are no records of this species ever occurring in some parts of the indicated distribution (e.g., Illinois; Jim Herkert, pers. comm.). This species is apparently extirpated in Connecticut and Ohio (where known from only one specimen). Elevational range extends to at least 700-800 meters in several states and to at least 1,125 meters in Kentucky (see Best and Jennings 1997).

M. leibii are one of the smallest bats in eastern North America. Total body length averages 83mmand tail length averages 36mm. Cranial dimensions average 13.6mm in length, 4.2mm in depth, and 5.1 mm for the maxillary tooth row. Their ears are less than 15mm long and their hind feet measure less than 8mm, a diagnostic feature of the small-footed Myotis. They have a wingspan of 210-250mm with a forearm length of 32.2mm and an average mass of 3.8g. (Best and Jennings 1997).

Eastern small footed bats have soft, thick, glossy fur with a yellowish tan to blackish brown colour and a golden sheen to the surface. The roots of the fur are black with pale brown tips which give the golden sheen. The ventral surface of their body is grey and their face, ears, wings and interfemoral membrane are black. Their ears reach or exceed the tip of their nose when laid forward and their tail extends beyond their naked interfemoral membrane. (Best and Jennings 1997).

Myotis leibii have a keeled calcar. Their skull is small and delicate and their braincase is flattened. The forehead slopes gradually upward from the rosturm, they lack the prominent forehead common to other Myotis species. The dental formula is 2/3I 1/1C 3/3P 3/3M for a total of 38 teeth. (Best & Jennings 1997).

Range mass: 3.2 to 5.5 g.

Average mass: 3.8 g.

Length: 8 cm

Weight: 9 grams

This bat differs from other sympatric bats by a) small size, < 8.5 cm, b) small hind foot, < 8 mm, c) black face, d) long-keeled calcar, and e) absence of a dark shoulder patch. It differs from M. lucifugus by having golden-tinted, almost yellowish fur and a shorter forearm. It differs from Pipistrellus subflavus by its lighter color, especially the light pinkish forearms, and lack of a keeled sternum (Godin 1977). The skull is much flatter than that of M. lucifugus, and the braincase is narrower; a sagittal crest may be present (Banfield 1974, Godin 1977, Schwartz and Schwartz 1981, Merritt 1987).

• Terrestrial

The Eastern small footed bat occurs in mountanous regions at elevations ranging from 240-1125m. They prefer eastern deciduous and coniferous forests and can roost in buildings, rock bluffs and turnpike tunnels during spring and summer months. (Best and Jennings 1997).

Hibernation sites are in mines and caves only. During hibernation they tend to hang near the entrance where there is low humidity. When the winters become harsh they retreat further back into the caves. They do not shift cave sites and it appears that they hibernate near their summer ranges. These bats have been found to hibernate horizontally under rock slabs possibly due to a more constant temperature and humidity under these conditions. (Barbour & Davis. 1969; Best & Jennings 1997; Davis 1955; Fenton 1972).

Terrestrial Biomes: forest

Comments: Habitat is mostly hilly or mountainous areas, in or near deciduous or evergreen forest, sometimes in mostly open farmland. In Pennsylvania, Mohr (1932) found this species mostly in heavy hemlock forests in the foothills of mountains that rise to 2,000 feet (600 meters). Unpublished data from the Kentucky Heritage Program indicate that summer roosts include caves, coal mines, buildings, and bridges over rivers (in expansion joints). Warm-season roosts include buildings, towers, hollow trees, spaces beneath the loose bark of trees, cliff crevices, and bridges. Tuttle (1964) reported two individuals found in April in Tennessee under a large flat rock at the edge of a quarry surrounded by woods and cow pastures. In Ontario, about 12 of these bats were found in July behind the door of a shed that was kept open (i.e., positioned against the wall) (Hitchcock 1955). They have been seen resting in limestone caves in West Virginia in spring and summer (Krutzsch 1966).

By far most records come from observations of bats hibernating in winter in caves and mine tunnels. Hibernation occurs in solution and fissure caves and mine tunnels (including coal, iron, copper, and talc mines). Situations near the entrance where the air is relatively cold and dry seem to be preferred (Barbour and Davis 1969), though sometimes deeper locations are used (Schwartz and Schwartz 1981). Roost sites often are deep in crevices, or under rocks on the cave floor, where the bats can be very difficult to find (Davis 1955, Krutzsch 1966, Martin et al. 1966). These bats are usually found singly or occasionally in small clusters, but many may be packed in a crevice; often they hang among other species (Marin et al. 1966). In tight places the body may be horizontal, even belly down. On cave walls, the forearms are somewhat extended rather than parallel to the body axis.

Dunn and Hall (1989) noted that 52% of Pennsylvania hibernacula were small caves of less than 150 m (500 feet) in length.

Like many other bat species, this one typically forages over ponds and streams.

Non-Migrant: No. All populations of this species make significant seasonal migrations.

Locally Migrant: Yes. At least some populations of this species make local extended movements (generally less than 200 km) at particular times of the year (e.g., to breeding or wintering grounds, to hibernation sites).

Locally Migrant: No. No populations of this species make annual migrations of over 200 km.

In Ontario, two were recovered in summer 16 and 19 km from the winter hibernation cave (Hitchcock 1955). Barbour and Davis (1969) reported: "In the Mammoth Cave region of Kentucky, M. LEIBII is fairly common in late summer in the flocks of migrating bats. The whereabouts of these individuals at other seasons in unknown."

Few details are known about the diet and feeding habits of M. leibii, although they appear to be insectivores. They have been seen to forage over water and land collecting insects while in flight. Favorite prey includes small insects such as flies, beetles, and moths which are caught on the wing. (Best & Jennings 1997; Wilson & Ruff 1999).

Comments: Probably the diet is almost exclusively flying insects; specific data are lacking. Feeding flights are relatively slow and fluttery and occur over ponds and streams and along roads.

Note: For many non-migratory species, occurrences are roughly equivalent to populations.

Estimated Number of Occurrences: 81 to >300

Comments: This species is most often detected during hibernation. In recent years, it has been counted at approximately 125 hibernacula. Recent surveys have greatly increased the number of localities above those known historically; the number of hibernacula may be significantly larger than currently known. Intensive cave and mine surveys have been undertaken in most states where the species occurs, but some sites probably remain unsearched in most states.

2500 - 100,000 individuals

Comments: The total count for all hibernacula is approximately 3,000 individuals, with roughly 60% of the total number from just two sites in New York. Some of the occurrences probably have not been surveyed completely, and some individuals are undoubtedly missed within some sites because they are hibernating in portions of mines or caves that cannot be reached or easily observed.

Significant diseases, parasites, and predators are not known. Mites (identification pending) have been found on the ears of an individual in South Carolina (Mary Strayer, pers. comm.). Related species are parasitized by trematodes, cestodes, chiggers, mites, fleas, and bat bugs (Cimex sp.). Possible predators include domestic cats, mink, raccoons, opossum, fish, frogs, snakes, and birds of prey. Most of these are known to prey occasionally on other Myotis species (Barbour and Davis 1969). This is one of the many mammals known to carry rabies (Constantine 1979).

Bats are often important components of cave ecosystems, providing the primary energy input in the food chain in the form of guano (Horst 1972, Poulson 1972).

Comments: Emerges from its daytime retreat shortly after sunset, while there is still some light. In the northeastern U.S., seldom enters hibernation caves before mid-November; departs by March, or possibly earlier in Vermont (Godin 1977).

Average lifespan

Status: captivity: 12 years.

There is a 1:1 sex ratio at birth. Mating occurs in autumn before hibernation. Sperm is stored in the female until fertilization in the early spring. Females have 2 mammae and give birth to a single young between late May and July. Females have been found in maternity colonies ranging from 12-20 individuals. The lifespan of M. leibii is thought to be between 6-12 years, however they have a low survival rate (0.757 for males and 0.421 for females). (Best & Jennings 1997; Hitchcock et al. 1984; Wilson & Ruff 1999).

During copulation males mount the female from behind, grasping the back of her neck with his teeth and further stabilizing her with his thumbs. The interfemoral membrane does not appear to be an obstacle during copulation due to the free movement of the male's penis. The females role is passive. There are no vocalizations during mating. (Best and Jennings 1997).

Range number of offspring: 1 to 1.

Key Reproductive Features: seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); viviparous ; sperm-storing

Average number of offspring: 1.

Mating behavior is similar to that of the little brown bat (Wimsatt 1945) and so, probably, are other facets of reproduction (Banfield 1974). Breeding may occur in the fall, with the sperm stored in the uterus over winter. Active hestation lasts probably two months, with a single offspring born annually, probably in early July (Merritt 1987).

Survival rates are significantly lower for females (42%) than for males (76%) (van Zyll de Jong 1985). One individual is reported to have lived 12 years (Hitchcock 1965). One summer colony included about 12 individuals (Hitchcock 1955). Colonies are usually small (< 15), though a few number in the hundreds.

Pennsylvania has listed M. leibii as threatened and at risk. The United States Fish and Wildlife Service has listed M. leibii as a candidate for protection under the Endangered Species Act. (Best & Jennings 1997)

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

US Federal List: no special status

CITES: no special status

IUCN Red List of Threatened Species: least concern

Canada

Rounded National Status Rank: N3 - Vulnerable

United States

Rounded National Status Rank: N3 - Vulnerable

Rounded Global Status Rank: G3 - Vulnerable

Reasons: Fairly widespread in southeastern Canada and eastern U.S., but very spotty in distribution and rarely found in large numbers; few high quality occurrences exist; the total numbers counted are very low in comparison to the total number of caves and mines surveyed; the total numbers counted and the number of occurrences are the primary consideration in the ranking; susceptability to disturbance is also an important factor.

Intrinsic Vulnerability: Highly to moderately vulnerable.

Environmental Specificity: Very narrow. Specialist or community with key requirements scarce.

Global Short Term Trend: Relatively stable (=10% change)

Comments: Most occurrences have been counted only within the past decade or two and are not revisited regularly, making the assessment of population trend difficult. Many biologists believe that this species is basically stable, having declined little in recent times, but that it is vulnerable, especially in its cave hibernacula. The population at one site in Arkansas has increased in recent years, probably due to reduced winter disturbance following the installation of a cave gate.

Global Long Term Trend: Increase of 10-25% to decline of 50%

Comments: This bat always has been considered to be relatively rare (Barbour and Davis 1969). Numbers are reduced in a few sites where older counts are available, and a few historical sites are apparently no longer occupied (e.g., see Hall 1979, but compare Dunn and Hall 1989), but whether these observation reflect declines or changes in distribution is unknown. In Vermont, Myotis leibii has been consistently found in very small numbers and often not detected at all during periodic surveys of various hibernacula dating back to 1934 (Trombulak et al. 2001).

Degree of Threat: B : Moderately threatened throughout its range, communities provide natural resources that when exploited alter the composition and structure of the community over the long-term, but are apparently recoverable

Comments: Some mines may be threatened by closure or collapse. Ceiling collapse may kill bats outright or, more significantly, alter cave microhabitat enough to make it unsuitable. A few cave occurrences are threatened or have been reduced in quality due to commercialization for tourism.

Threats to summer sites are unknown, but are likely to be moderate due to alteration of riparian habitats. Conversion of forested habitats to agricultural and residential uses has decreased the amount of preferred habitat in some areas, but the bats do make use of bridges and various other non-natural roost sites.

Improper gating of caves to protect bats may result in site abandonment. For example, the large colony of M. leibii at Fourth Chute Cave, Quebec, was driven out by blocking the flow of cold air (Mohr 1972). In contrast, gating of cave entrances in other locations (e.g., Aitkin Cave, Pennsylvania) has led to increases in M. leibii populations.

With its small numbers and spotty distribution, isolated colonies of M. leibii are particularly vulnerable to extirpation by chance events, especially when concentrated during winter months. On the other hand, in contrast to certain other bats that assemble in vast numbers in relatively few sites, the population of M. leibii as a whole is not vulnerable to localized events.

This bat tends to hibernate near cave entrances; hence it may be vulnerable to freezing in abnormally severe winters.

THREATS TO BATS IN GENERAL: Perhaps the most serious threat to cave-dwelling bats is human disturbance during hibernation. Very low levels of noise, light, and heat from lanterns are sufficient to awaken hibernating bats, which then expend energy moving about and deplete critical reserves of body fat. When disturbance is repeated, bats (especially juveniles) are likely to perish. "By the end of the winter energy reserves may be insufficient to meet the demands of the first feeding forays, when emerging insects may be scattered and scarce, or the bats may be too weak to make long flights to their summer territories" (Mohr 1976). Such disturbance is equally lethal, whether caused by vandals, well-meaning spelunkers, or bat researchers.

Intentional killing of bats in caves by clubbing, stoning, burning, bombing, etc. has been a significant cause of mortality. Documented examples are numerous (Greenhall 1973, Harvey 1976, Tuttle 1979, and others). Bats are sometimes exterminated from commercial caves, or if not, leave or move to suboptimal habitats due to increased disturbance.

Poisoning by pesticides, heavy metals, and other environmental contaminants has been and may remain a significant threat (Clark 1988). Destruction of roost and foraging habitat by reservoir inundation, strip mining (especially limestone), deforestation, drainage of wetlands, development, etc., and pollution or siltation of waterways with consequent decline in insect production are additional potential adverse impacts (Tuttle 1979). Hundreds of thousands of bats have been destroyed by natural flooding of caves (Hall 1962, Brady 1979).

Restoration Potential: Restoration to formerly occupied caves may be possible if conditions of temperature, air flow, moisture, and access remain unchanged, but there is little guiding information. Mohr (1976) thought this an unlikely option: "Attempts to transfer bat populations from houses to tunnels in Pennsylvania (Mohr 1942) were totally unsuccessful and recent evidence of the extreme loyalty--philopatry--of bats to their accustomed caves (Tuttle 1975) leaves little hope for reestablishing deserted hibernating sites." There is evidence of Indiana bats (Myotis sodalis) and big-eared bats (CORYNORHINUS/PLECOTUS spp.) returning to abandoned caves after gating or fencing (Craig Stihler, pers. comm.) In the case of gray bats (Myotis grisescens), Tuttle (1979) thought they might return to previously used caves, but that "any cave not already used by gray bats, however, should be assumed to be unsuitable for future use. Such caves probably do not provide essential temperature or roosting conditions, are too distant from acceptable foraging or hibernating sites, or are too vulnerable to predation or flooding." He thought that all suitable caves were probably occupied long before the arrival of man. Myotis leibii (among others) has found new habitat in certain mine tunnels, but little is known about the specific microhabitat conditions required.

Preserve Selection and Design Considerations: Preserves should include sufficient buffer area above and around the hibernaculum cave to protect it from disturbances that might alter water and air flow, temperature, and humidity. Adjacent to summer roosts and maternity sites (in the unlikely event that they are known), sufficient foraging area (probably along a stream) should be protected from deforestation, contamination with pesticides, and other major impacts. Too little is known about this species' ecology to make more specific recommendations.

Management Requirements: With the present scarcity of pertinent information, management can focus only on protection of hibernation caves summer roosts. The locations of caves, especially in remote areas, should not be publicized, and trails to entrances should be eliminated. Depending on the situation and potential for irresponsible traffic or vandalism, it may be useful to construct a fence or steel gate and/or place warning/interpretive signs. Hibernacula should be closed from November through March.

Public education as to the true nature and value of bats could avoid many problems and is urgently needed. Staff of national and state forests and parks, wildlife agencies, agricultural extension agents, and others in similar positions should be educated first so they can enlighten the public.

Foraging areas must be identified and protected from pesticides and other contaminants that may poison bats directly as well as destroy their food sources. Analysis of guano may be a convenient way of checking colonies for contamination (Clark 1988), in the rare cases of where summer colonies are known and there is sufficient accumulation. Bats for diagnostic study are generally frozen immediately. Consult a U.S. Fish and Wildlife Service Environmental Contaminant Field Specialist in cases of suspected poisoning or disease.

Logging above and around bat caves should be limited; deforestation can alter cave temperature, humidity, and air and water flow. Blasting in limestone mines or winter logging in the immediate vicinity of caves could cause arousal of hibernating bats. Some forest cover around the cave entrance and foraging area may offer significant protection from predators and periods of exceptionally cold spring weather (Tuttle 1979). Some Myotis species seem to require wooded corridors from roosts to feeding areas (Craig Stihler, pers. comm.).

Harvey (1981) offered cave protection suggestions: "Where gates are necessary, they should be used with extreme care to avoid detrimental effects to bats and cave microclimates. Each gate must be designed specifically for the cave to be protected, considering number of bats, type of colony, air flow, and entrance size and configuration. They should not be used at summer caves unless adequate free flight space can be provided above. Gates should be vertical (not horizontal or slanting) and should not be placed in entrances smaller than 1.8 m (6 ft) in diameter. Care must be taken so that gates do not restrict normal air flow. They should be constructed of welded steel bars 1.9 cm (3/4 inch) to 2.5 cm (1 inch) in diameter and of sufficient hardness to be invulnerable to bolt cutters. Free ends of bars should be grouted into solid rock. If a concrete footing is necessary, it should not extend above original ground level. Access openings should be constructed with durable hinges, hasps, and locks. Openings through which bats are expected to fly should be 15.2 cm (6 inches) vertically and 61 cm (24 inches) horizontally. In situations where vandalism is likely, a weak-link design may be employed. The lock, hasp, or some other easily replaceable part should be relatively weak so that vandals, if determined, can break in without doing excessive damage to the gate." [Modern gates constructed with 4 in. (10 cm) angle iron may make this unnecessary; also, latest designs minimize the number of vertical members, which seem to disturb bats (Craig Stihler, pers. comm.).]

"At some caves, especially those with sinkhole entrances, a simple vertical gate is impractical. In these situations a 'cage' or 'box' type gate may be necessary so that bats can fly up into the 'box' and then out through vertical openings. If gates are used at summer caves, they should be 'half-gates.' Such a gate is practical only in large cave entrances, where it can extend part way to the ceiling and still allow adequate space (at least 1 m, ideally more) above the gate through which bats can fly" (Harvey 1981). ["Half-gates" are probably only necessary for caves used also by gray bats (Craig Stihler, pers. comm.).] After gates are placed at cave entrances, bat activity should be monitored to assure that gates are not having detrimental effects on bat populations. Gates should be inspected periodically and often so that necessary repairs can be made promptly." (Harvey 1981).

"Fences do not afford as much protection as steel gates and are easier to cut or climb. Nevertheless, circumstances at some caves make fences more practical, especially in situations where gates might result in abandonment of the cave by bats. Chainlink, barbed-wire-topped fences with posts set in concrete are recommended. Barbed-wire should not extend into flight space required by bats" (Harvey 1981). [The bottoms of fences must also be secured; people going underneath are a problem (Craig Stihler, pers. comm.).]

"At caves which are infrequently visited, or which can be easily observed by the landowner, a sign alone may be adequate to prevent disturbance. In some circumstances, a sign might attract unnecessary attention to a cave, in which case the management agency might place a sign inside the cave, or not use a sign." (Harvey 1981).

"Signs should be of durable construction and fixed solidly in place to minimize vandalism, and should be placed so as to not interfere with bat flight patterns or air flow. They should be located where potential violators will see them, and should be placed behind the gate or fence if such a structure has been erected. Wording may vary from cave to cave, but signs should contain both a warning and interpretive message. At many caves it may be permissible to permit entry during times of the year when bats are not present. A sign containing that message, plus information on how to obtain a key to gated or fenced caves, might discourage vandals and encourage the cooperation of spelunkers. Where a cave is located in a public use area, the management agency may wish to use a more elaborate sign with a more detailed interpretive message" (Harvey 1981).

White and Seginak (1987) also discussed gate designs for bat caves.

Management Research Needs: Very little is known about the ecology, reproduction, and life history of this species. Especially needed is knowledge of critical habitat requirements, including hibernacula, summer roost and maternity sites, and foraging areas, food sources and impacts on them, and reproductive biology. Movements between hibernacula and summer sites need to be investigated.

The potential for translocation of colonies threatened by habitat destruction needs study (Mohr 1976).

There is still a need for new monitoring methods that are accurate and minimize impact on the bats.

Global Protection: Few to several (1-12) occurrences appropriately protected and managed

Comments: There is a strong conservation easement held by The Nature Conservancy (TNC) on the second largest occurrence. TNC has a management agreement giving limited protection to the largest occurrence and one occurrence with apparently good viability is on National Park Service land in Arkansas. Several occurrences with fair to poor estimated viability in various states are either owned by TNC or are on Federal land.

Needs: All A and B quality occurrences should be fully protected along with as many C occurrences as possible. Protection should include the ground surface above and adjacent to caves and mines and may need to include mineral rights for some mines. Protection, whether by ownership, easement or management agreement must include some means of limiting winter disturbance.

Myotis leibii are known to commonly roost in human structures and therfore may be considered a nuisance by some. (Best & Jennings 1997).

As insectivores, M. leibii may help control insect pests, including mosquitos, which they have been observed catching in the lab. However, because of their small population size there may be little or no noticeable effect. (Best and Jennings 1997).

Stewardship Overview: The eastern small-footed myotis is most vulnerable during hibernation. The caves and mines that serve as significant hibernacula must be protected from disturbance November through March. If necessary, the entrance should be gated. Foraging areas (mostly streams and ponds) should be protected from pesticides and anything else that might adversely affect production of the bat's insect food. Forest should not be eliminated above or around hibernacula, nor around foraging areas. Public education about the value of bats is necessary in the long-term.

Comments: Formerly, M. ciliolabrum was included as a subspecies of M. leibii (or M. subulatus). Based chiefly on cranial measurements, Van Zyll de Jong (1984) recognized western populations of what had been known as M. subulatus as a species (M. ciliolabrum) distinct from eastern populations, for which the appropriate name is M. leibii. Electrophoretic data support the conclusion that the two taxa are specifically distinct (Herd 1987). Wilson and Ruff (1999) and Baker et al. (2003) regarded M. ciliolabrum and M. leibii as separate species. Koopman (in Wilson and Reeder 1993) did not recognize M. ciliolabrum as a species distinct from M. leibii, but Simmons (in Wilson and Reeder 2005) did. See also Miller and Allen (1928), Glass and Baker (1968), and Hall (1981) for nomenclatural information. Monotypic (Best and Jennings 1997).

In a phylogenetic study based on mtDNA data, M. leibii was included within clades containing both M. californicus and M. ciliolabrum (Rodriguez and Ammerman 2004). Further data from M. leibii are necessary to validate its phylogenetic relationship to M. ciliolabrum and M. californicus (Rodriguez and Ammerman 2004). Comparisons among outgroups (M. yumanensis, M. lucifugus, and M. evotis) found sufficient support for specific status of M. leibii, but sequence divergence between M. evotis and the leibii group was small (2.9%) and within the intraspecific range. Further sampling of M. evotis is necessary to establish the level of divergence between M. evotis, as well as other long-eared Myotis, and the leibii group (Rodriguez and Ammerman 2004).