Leptonycteris nivalis - (Saussure, 1860)
Mexican Long-nosed Bat
Taxonomic Status: Accepted
Related ITIS Name(s): Leptonycteris nivalis (Saussure, 1860) (TSN 180068)
Spanish Common Names: Un Murciélago
Unique Identifier: ELEMENT_GLOBAL.2.104259
Element Code: AMACB03010
Informal Taxonomy: Animals, Vertebrates - Mammals - Bats
 
Kingdom Phylum Class Order Family Genus
Animalia Craniata Mammalia Chiroptera Phyllostomidae Leptonycteris
Genus Size: B - Very small genus (2-5 species)
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Concept Reference
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Concept Reference: 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/.
Concept Reference Code: B93WIL01NAUS
Name Used in Concept Reference: Leptonycteris nivalis
Taxonomic Comments: Prior to 1962 (Davis and Carter 1962), specimens of Leptonycteris yerbabuenae were reported as L. nivalis.
Conservation Status
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NatureServe Status

Global Status: G3
Global Status Last Reviewed: 19Mar2015
Global Status Last Changed: 19Mar2015
Ranking Methodology Used: Ranked by calculator
Rounded Global Status: G3 - Vulnerable
Reasons: Widely distributed but relatively rare in Mexico; also occurs in a few areas in southwestern Texas and southwestern New Mexico; recent population size and trend are not well documented; known primary threats include disturbance of cave roosts by humans and loss/degradation of foraging habitat/food resources (e.g., wild agave) as a result of human activities.
Nation: United States
National Status: N2 (16May2014)

U.S. & Canada State/Province Status
United States New Mexico (S2), Texas (S1)

Other Statuses

U.S. Endangered Species Act (USESA): LE: Listed endangered (30Sep1988)
U.S. Fish & Wildlife Service Lead Region: R2 - Southwest
IUCN Red List Category: EN - Endangered

NatureServe Global Conservation Status Factors

Range Extent: 200,000-2,500,000 square km (about 80,000-1,000,000 square miles)
Range Extent Comments: Range includes northern and central Mexico, southwestern Texas, and southwestern New Mexico, generally at elevations of about 500 to 3,000+ meters. Most occurrences in Mexico are at elevations of 1,000-2,200 meters, but this bat been captured at an elevation of 3,780 meters (see Arita 1991), and the type specimen reportedly was caught near snow line at 17,816 feet (5,747 meters) on Mt. Orizaba, in Veracruz, Mexico (USFWS 1994). In Texas, the species has been captured in Big Bend National Park (Brewster County) and the Chinati Mountains (Presidio County); Emory Peak Cave in the Chisos Mountains (elevation 2,290 meters) hosts the only known roosting population in Texas (Ammerman et al. 2012). Two specimens of Leptonycteris taken in Hidalgo County, New Mexico (in 1963 and 1967), were determined to be L. nivalis. The presence of this species in New Mexico was reconfirmed in Hidalgo County in 1992 (Hoyt et al. 1994). Populations exist in the Animas and Big Hatchet mountain in New Mexico (P. Cryan, pers. comm., cited by Ammerman et al. 2012). The range has been described as extending into Guatemala and adjacent southern Mexico (Hensley and Wilkins 1988; Simmons, in Wilson and Reeder 2005), but specimens collected from those areas were assigned to L. yerbabuenae by Arita and Humphrey (1988) and Arita (1991). Simmons (in Wilson and Reeder 2005) described the range of L. nivalis as including southeastern Arizona, but no actual records for Arizona are known.

Area of Occupancy:  
Area of Occupancy Comments: Suitable roosts may be a limiting factor; area of occupancy is quite small if based on the known area (number of 2 km x 2 km grid cells) occupied by roosting bats. In contrast, according to modeling algorithms (GARP and Maxent), the estimated area of suitable foraging habitat is up to 709,525 square kilometers (Emma Gomez-Ruiz and Thomas E. Lacher, unpublished data), 

Number of Occurrences: 21 - 80
Number of Occurrences Comments: The number of occurrences or subpopulations has not been determined using standardized/meaningful criteria; if based on regularly occupied roost sites, the number probably exceeds 20 and may not exceed 80. Arita and Humphrey (1988) and Arita (1991) mapped 43 confirmed collection sites for L. nivalis in Mexico, though not all of these necessarily represent roost sites. Just a few roosting sites exist in the United States, but there could be additional sites that have not yet been detected.

Population Size: 10,000 - 1,000,000 individuals
Population Size Comments: Total adult population size is unknown but presumably exceeds 10,000 and may exceed 100,000, based on the extensive range and the periodic presence of several thousand individuals in the single roost in Texas. This species is widespread but "scarce" in most of its range (Arita 1993, Arita and Santos-del-Prado 1999). In Mexico, the species forms cave colonies of fewer than 100 individuals or 100-10,000 individuals (Arita 1993). Moreno-Valdez et al. (2004) recorded a population of up to a few thousand individuals at El Infierno Cave, Nuevo Leon, Mexico, in the late 1990s.

Roost at Emory Peak Cave (Texas): 10,650 bats in 1967, 5,000 in 1968, 3,900 in 1969, 0 in 1970, 8025 in 1971, 1000 in 1983, 4,942 -5,990 in 1988, 5,000+ in 1991, 0 in 1992, and 2,859 in 1993 (Matthews and Moseley 1990, see USFWS 1994). Large annual fluctuation may be in part due to some counts being made before or after the main period of bat occupation of the cave (Cockrum and Petryszyn 1991), or it may be an artifact of some bats being present but not detected (Ammerman et al. 2009), or it may reflect the movements of bats in response to available food resources (Moreno-Valdez et al. 2004, Ammerman and Tabor (2008).

Number of Occurrences with Good Viability/Integrity: Some (13-40)
Viability/Integrity Comments: The number of roost-based occurrences or subpopulations with good viability is uncertain but likely there are not very many. Several caves in central Mexico were known to contain considerable numbers in the past but now contain only small colonies or none at all (see USFWS 1994).

Overall Threat Impact: High - medium
Overall Threat Impact Comments: Primary threats include disturbance of roosts, loss of food sources through clearing of land for agriculture and human exploitation of agaves (e.g., for production of alcoholic beverages), and direct killing by humans (USFWS 1987, USFWS 1988, USFWS 1994). Other threats may include negative effects of ingestion of pesticides applied to plants, competition for roosts and nectar, natural catastrophes, disease, and predation; however, these are not believed to be major limiting factors, though some of them could become significant for populations reduced to small size by other factors (see USFWS 1994).

These bats are sensitive to disturbance in their roosting sites (they often quickly take flight upon human entry; Wilson 1985, Wilson et al. 1985) and, in general, roosting caves are becoming increasingly subject to human destruction and disturbance. Human disturbance and destruction of roost sites is a common occurrence in Mexico. The availability of roost sites free from disturbance may be a significant limiting factor.

A major problem for bats all over Mexico is that uninformed citizens frequently destroy all bats in a roost, believing them to be vampire bats (USFWS 1994).

The only known mating site for the species, Cueva del Diablo located in Tepoztlan, Morelos, Mexico, is threatened by residential development for tourism (Emma P. Gomez-Ruiz and Thomas E. Lacher, pers. comm., 2015).

Foraging habitat can be degraded or destroyed by harvesting of agave (an important food resource). It has been estimated that bootleg mescal makers are eliminating between 500,000 and 1,200,000 wild paniculate agave a year in Sonora alone. Agave plant parts are harvested just before they bloom; this prevents flowering and can prevent reproduction by the affected plant (an agave plant grows for 10 to 20 years, flowers only once, then dies). However, there are few places in Sonora or elsewhere in Mexico where wild Agave harvesting has eliminated a significant percentage of nectar-producing genets, and plants harvested by indigenous people generally produce vegetative offshoots that may eventually produce flowers (Nabhan and Fleming 1993). Populations of unharvested plants persist in many areas that are not easily accessible to humans. The negative impact of agave harvest probably is not as great as was previously believed. On the other hand, reduced pollination resulting from decreased bat populations may eventually lead to a reduction in agave distribution and abundance.

Preliminary data suggest that Agave is the main food source for L. nivalis in Coahuila and Nuevo Leon (Emma Gomez and Thomas Lacher, unpublished data). Human exploitation of wild agaves in the northern range needs to be evaluated to quantify this potential threat. Agaves are used extracting the nectar, and during intense drought periods the plants are harvested to feed cattle.

Some foraging habitat has been degraded or destroyed by expansion of agriculture and other land uses. Large areas of both the Sierra Madre Oriental and Sierra Madre Occidental and the Mexican Plateau have been converted to agriculture or rangeland.

Land clearing is a threat in northern Mexico, especially with recent plans for shale gas extraction and wind farm development (Emma P. Gomez-Ruiz and Thomas E. Lacher, pers. comm., 2015). Wind farming being developed in Coahuila and Nuevo Leon represents a risk, especially in areas where L. nivalis is known to occur during the summer migration. Evidence suggests that mainly pregnant females migrate to the North (maternity caves have only been reported in the North); wind turbine fatalities of pregnant females may pose a high risk to population stability (a better understanding of this risk is needed) (Emma P. Gomez-Ruiz and Thomas E. Lacher, pers. comm., 2015).

Changes in precipitation levels and timing have an effect on blooming events of Agave spp. (Emma P. Gomez-Ruiz and Thomas E. Lacher, pers. comm., 2015), so ongoing climate change potentially will affect this species.

This species may experience predation from owls, hawks, snakes, and mammals, but natural levels of predation likely are inconsequential to the overall status of the species. However, increased populations of domestic and feral cats and other predators near human habitations may affect the survival of colonies, particularly maternity colonies (USFWS 1994).

Short-term Trend: Unknown
Short-term Trend Comments: Trend over the past 10 years or three generations is uncertain. At Cueva del Diablo in Morelos, Medellín (2003) reported an increase from an estimated 5,000 in 1996 to 8,000-10,000 in the winter of 2001-2002.

Long-term Trend: Decline of 30-70%
Long-term Trend Comments: Long-term trend is unclear. Extent of occurrence and area of occupancy probably have not changed much, but the number of occurrences or subpopulations and population size may have dramatically decreased in some locations during the last three decades. Wilson (1985) found that this species was either completely absent or present in reduced numbers in known roosts. The number of bats found represented only a fraction of the total reported in previous studies. For example, in an abandoned mine in Nuevo Leon, Mexico, where an estimated population of 10,000 was observed in 1938, no individuals of L. nivalis were found in 1983 (Wilson 1985). Another mine in Nuevo Leon had a ceiling covered with newborn bats in 1967, but only one bat was found in 1983. A few other roosts had reduced numbers of bats compared to findings during previous surveys. These changes could indicate a decline in the overall population, but they might reflect movement of bats among different roosting sites in different years, or they could result from seasonal changes in bat distribution (survey dates varied).

A colony of L. nivalis in Morelos, Mexico, increased from an estimated 5,000 in 1996 to 8,000-10,000 in 2001-2002 (Medellín 2003).

Abundance at Emory Peak Cave in Texas fluctuates widely from year to year (0 to 10,000+ individuals). Reasons for the fluctuations are not completely understood, but they apparently reflect annual variations in regional food resources (number of flowering agave plants) (USFWS 1994, Ammerman and Tabor 2008); a similar pattern has been observed at a cave in Nuevo Leon, Mexico (Moreno-Valdez et al. 2004). Historical count data for Emory Peak Cave may not be completely reliable; bats present in the cave may go undetected (Ammerman et al. 2009, 2012).

Ammerman et al. (2012) noted a lack of consistency among various trend estimates or indications for this species (Wilson 1985, Arita and Humphrey 1988, Cockrum and Petryszyn 1991, USFWS 1994, Medellín 2003, Ammerman and Tabor 2008, Ammerman et al. 2009) Despite some inconsistency, most authors have concluded that the species is declining in abundance, though the degree of decline is highly uncertain. However, better population data based on improved monitoring methods are needed before a reliable trend determination can be made (Ammerman et al. 2012).

Other NatureServe Conservation Status Information

Inventory Needs: Further regular surveys are needed to determine important roost sites, abundance, and trends.

Protection Needs: Conservation of this bat will require maintenance of relatively large areas of wild agave (Moreno-Valdez et al. 2004). Identification and protection of currently and formerly occupied roost sites and protection of foraging habitat in at least several areas throughout the range are important conservation needs.

Distribution
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Global Range: (200,000-2,500,000 square km (about 80,000-1,000,000 square miles)) Range includes northern and central Mexico, southwestern Texas, and southwestern New Mexico, generally at elevations of about 500 to 3,000+ meters. Most occurrences in Mexico are at elevations of 1,000-2,200 meters, but this bat been captured at an elevation of 3,780 meters (see Arita 1991), and the type specimen reportedly was caught near snow line at 17,816 feet (5,747 meters) on Mt. Orizaba, in Veracruz, Mexico (USFWS 1994). In Texas, the species has been captured in Big Bend National Park (Brewster County) and the Chinati Mountains (Presidio County); Emory Peak Cave in the Chisos Mountains (elevation 2,290 meters) hosts the only known roosting population in Texas (Ammerman et al. 2012). Two specimens of Leptonycteris taken in Hidalgo County, New Mexico (in 1963 and 1967), were determined to be L. nivalis. The presence of this species in New Mexico was reconfirmed in Hidalgo County in 1992 (Hoyt et al. 1994). Populations exist in the Animas and Big Hatchet mountain in New Mexico (P. Cryan, pers. comm., cited by Ammerman et al. 2012). The range has been described as extending into Guatemala and adjacent southern Mexico (Hensley and Wilkins 1988; Simmons, in Wilson and Reeder 2005), but specimens collected from those areas were assigned to L. yerbabuenae by Arita and Humphrey (1988) and Arita (1991). Simmons (in Wilson and Reeder 2005) described the range of L. nivalis as including southeastern Arizona, but no actual records for Arizona are known.

U.S. States and Canadian Provinces
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Endemism: occurs (regularly, as a native taxon) in multiple nations

U.S. & Canada State/Province Distribution
United States NM, TX

Range Map
Note: Range depicted for New World only. The scale of the maps may cause narrow coastal ranges or ranges on small islands not to appear. Not all vagrant or small disjunct occurrences are depicted. For migratory birds, some individuals occur outside of the passage migrant range depicted. For information on how to obtain shapefiles of species ranges see our Species Mapping pages at www.natureserve.org/conservation-tools/data-maps-tools.

Range Map Compilers: Sechrest, 2002


U.S. Distribution by County Help
State County Name (FIPS Code)
NM Hidalgo (35023)
TX Brewster (48043), Presidio (48377)*
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
13 Cibolo-Red Light (13040201)+*, Terlingua (13040204)+, Big Bend (13040205)+
15 Animas Valley (15040003)+
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
Ecology & Life History
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Basic Description: A leaf-nosed bat.
General Description: This grayish brown bat has a leaflike nose projection and no tail; forearm 55-60 mm (Whitaker 1996).
Diagnostic Characteristics: Leptonycteris curasoae is usually smaller, with a shorter forearm (51-56 mm) (Whitaker 1996). Choeronycteris mexicana has a small tail.
Reproduction Comments: Litter size normally is 1. Young are born apparently in spring (April-June), primarily in Mexico before females arrive in Texas, though pregnant females have been captured in Texas in late April (Brown 2008, Ammerman et al. 2012). In Texas, lactating females have been observed in June-July, flying juveniles in late June. Young are weaned in July or August. These bats are highly colonial.
Ecology Comments: These bats are effective pollinators of cacti and agave; the plants are dependent on bats for sexual reproduction.
Habitat Type: Terrestrial
Non-Migrant: Y
Locally Migrant: Y
Long Distance Migrant: Y
Mobility and Migration Comments: This species is migratory in the northern portion of its range (Wilson et al. 1985, Schmidly 1991, Ammerman et al. 2012), but movements are not well-known. Seasonal movements likely correspond with food availability. It has been recorded in the United States from June to August (Ammerman et al. 2012). Most northward migrants in Texas are females, but in New Mexico the sex ratio is more balanced (Hoyt et al. 2004; P. Cryan, pers. comm., cited by Ammerman et al. 2012).
Palustrine Habitat(s): Riparian
Terrestrial Habitat(s): Desert, Woodland - Mixed
Subterranean Habitat(s): Subterrestrial
Habitat Comments: Habitats include desert scrub, open conifer-oak woodlands, and pine forests in the Upper Sonoran and Transition Life Zones; generally arid areas where agave plants are present (USFWS 1994). Colonies roost in caves (or similar mines and tunnels), sometimes in culverts, hollow trees, or unused buildings. Roosting habitat requirements are not well known.
Adult Food Habits: Frugivore, Nectarivore
Immature Food Habits: Frugivore, Nectarivore
Food Comments: Diet includes mainly nectar and pollen of at least 21 plant species representing 10 plant families (Sánchez and Medellín 2007). In the northern part of the range, the bats often feed at the flowers of cacti and paniculate agaves. In Texas; nectar of mescal and Chisos agave flowers probably are the main food (Schmidly 1977). The diet may include insects associated with flowers, and probably some fruits, especially in the southern part of the range.
Adult Phenology: Nocturnal
Immature Phenology: Nocturnal
Phenology Comments: Activity occurs throughout the year. Emergence to feed occurs relatively late in the evening.
Colonial Breeder: Y
Length: 9 centimeters
Weight: 21 grams
Economic Attributes Not yet assessed
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Management Summary
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Stewardship Overview: Conservation of this bat likely will require maintenance of relatively large areas of wild Agave (Moreno-Valdez et al. 2004). Planting of agave has been initiated in some areas of northern Mexico. Public education is needed to reduce the level of human disturbance and destruction of roost sites in Mexico. Populations need to be defined, then several of them throughout the range need to be adequately protected. See recovery plan (USFWS 1994).
Biological Research Needs: Populations need to be defined, and movement patterns need to be better understood. Accurate censusing methods need to be developed and implemented.

The impacts of wild agave extraction/use by humans should be evaluated, particularly in the northern range where agave is the bats? main food source. A long-term monitoring program to document blooming events in agaves needs to be developed throughout the region; this could be implemented through a citizen science program. The potential impacts of wind farm development on L. nivalis habitat and on migrating bats need to be assessed. An economic evaluation of the pollination service provided by L. nivalis would be useful. Source: Emma P. Gomez-Ruiz and Thomas E. Lacher (pers. comm., 2015).
 

Population/Occurrence Delineation
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Group Name: Phyllostomid Bats

Use Class: Bachelor colony
Subtype(s): Roost Site, Foraging Area
Minimum Criteria for an Occurrence: An area occupied either historically or at present by a persisting or recurring bachelor male population. Identification evidence minimally includes collection or reliable observation and detailed documentation of one or more individuals.
Mapping Guidance: The EO includes both the colony site and the associated foraging areas. If separate, the colony site and foraging areas are bounded by separate polygons; that is, areas over which the bats simply commute to and from foraging areas and the colony are not included in the EO.
Separation Barriers: None.
Separation Distance for Unsuitable Habitat: 5 km
Separation Distance for Suitable Habitat: 5 km
Separation Justification: Movement characteristics of these highly mobile bats would suggest separation distance of a few to many tens of kilometers. However, this would result in occurrences of unwieldy spatial scope. It is impractical to attempt to delineate occurrences on the basis of discrete populations. Instead, the assigned separation distance is intended to generate occurrences that consist of spatially proximate roost sites and capture locations. However, include in the same occurrence (1) any roost sites between which individuals are known to move, regardless of how far apart they are, and (2) known significant foraging areas of occurrences that are based on roost sites.
Date: 24Mar2004
Author: Hammerson, G.

Use Class: Maternity colony
Subtype(s): Foraging Area, Colony Site
Minimum Criteria for an Occurrence: An area occupied either historically or at present by a persisting or recurring population of breeding females and their dependent young during lactation. Includes collections or mist net captures of pregnant or nursing females away from roost sites even if the actual roost site(s) are not known. Identification evidence minimally includes collection or reliable observation and detailed documentation of one or more pregnant or nursing females.
Mapping Guidance: The EO includes both the colony site and the associated foraging areas. If separate, the colony site and foraging areas are bounded by separate polygons; that is, areas over which the bats simply commute to and from foraging areas and the colony are not included in the EO.
Separation Barriers: None.
Separation Distance for Unsuitable Habitat: 5 km
Separation Distance for Suitable Habitat: 5 km
Separation Justification: Movement characteristics of these highly mobile bats (see following) would suggest separation distance of a few to many tens of kilometers. However, this would result in occurrences of unwieldy spatial scope. It is impractical to attempt to delineate occurrences on the basis of discrete populations. Instead, the assigned separation distance is intended to generate occurrences that consist of spatially proximate roost sites and capture locations. However, include in the same occurrence (1) any roost sites between which significant numbers of individuals are known to move, regardless of how far apart they are, and (2) known significant foraging areas of occurrences that are based on roost sites.

Radio-telemetry studies indicate that Macrotus in the California desert forage within 10 km of their roost (Brown, in Wilson and Ruff 1999). Female Leptonycteris curasoae forage over large distances; actual distance dependent on separation of appropriate roosting cave and appropriate feeding habitat. In nursing season (early summer), mean flight distance to foraging area about 14 kilometers at the Bluebird Cave in Arizona, but will fly up to 50 to 100 kilometers from their day roost (USFWS 1995, Westland Resources 2000). Normally fly up to 30 kilometers from their day roost (T. Fleming, pers. comm.).

Inferred Minimum Extent of Habitat Use (when actual extent is unknown): 10 km
Inferred Minimum Extent Justification: IE distance is based on the foraging range of MACROTUS CALIFORNICUS (Brown, in Wilson and Ruff 1999). Mean flight distance of LEPTONYCTERIS CURASOAE to foraging area is farther (USFWS 1995).
Date: 29Mar2004
Author: Cannings, S., and G. Hammerson

Use Class: Nonbreeding
Subtype(s): Foraging Area, Roost Site
Minimum Criteria for an Occurrence: An area occupied either historically or at present by a persisting or recurring population of nonbreeding individuals. Includes mist net captures or other detections away from roost sites obtained during the nonbreeding season even if the actual roost site(s) are not known. Identification evidence minimally includes collection or reliable observation and detailed documentation of one or more individuals.
Separation Barriers: None.
Separation Distance for Unsuitable Habitat: 5 km
Separation Distance for Suitable Habitat: 5 km
Separation Justification: Movement characteristics of these highly mobile bats would suggest separation distance of a few to many tens of kilometers. However, this would result in occurrences of unwieldy spatial scope. It is impractical to attempt to delineate occurrences on the basis of discrete populations. Instead, the assigned separation distance is intended to generate occurrences that consist of spatially proximate roost sites and capture locations. However, include in the same occurrence (1) any roost sites between which individuals are known to move, regardless of how far apart they are, and (2) known significant foraging areas of occurrences that are based on roost sites.
Inferred Minimum Extent of Habitat Use (when actual extent is unknown): 10 km
Inferred Minimum Extent Justification: Foraging range of MACROTUS CALIFORNICUS (Brown, in Wilson and Ruff 1999). Mean flight distance of LEPTONYCTERIS CURASOAE to foraging area is slightly farther, 14 kilometers (USFWS 1995).
Date: 29Jan2002
Author: Cannings, S.
Notes: Includes bats in the genera MACROTUS, CHOERONYCTERIS, and LEPTONYCTERIS.
Population/Occurrence Viability
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U.S. Invasive Species Impact Rank (I-Rank) Not yet assessed
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Authors/Contributors
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NatureServe Conservation Status Factors Edition Date: 30Mar2015
NatureServe Conservation Status Factors Author: Hammerson, G.
Element Ecology & Life History Edition Date: 02Apr2015
Element Ecology & Life History Author(s): Hammerson, G.

Zoological data developed by NatureServe and its network of natural heritage programs (see Local Programs) and other contributors and cooperators (see Sources).

References
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  • Ammerman, L. K., C. L. Hice, and D. J. Schmidly. 2012. Bats of Texas. Texas A & M University Press, College Station, Texas. xvi + 305 pp.

  • Ammerman, L. K., M. McDonough, N. I. Hristov, and T. H. Kunz. 2009. Census of the endangered Mexican long-nosed bat, Leptonycteris nivalis, in Texas. Endangered Species Research 8:87-92.

  • Ammerman, L. K., and R. Tabor. 2008. Monitoring the colony size and population fluctuations of the endangered Mexican long-nosed bat in Big Bend National Park using thermal imaging. Division of Science and Resource Management, Big Bend National Park. 8 pp.

  • Arita, H. T. 1991. Spatial segregation in long-nosed bats, Leptonycteris nivalis and Leptonycteris curasoae, in Mexico. Journal of Mammalogy 72:706-714.

  • Arita, H. T. 1993. Conservation biology of the cave bats in Mexico. Journal of Mammalogy 74:693-702.

  • Arita, H. T., and K. Santos-del-Prado. 1999. Conservation biology of nectar-feeding bats in Mexico. Journal of Mammalogy 80:31-41.

  • Arita, H. T., and S. R. Humphrey. 1988. Revisión taxonómica de los murciélagos magueyeros del género Leptonycteris (Chiroptera: Phyllostomidae). Acta Zoologica Mexicana (n.s.) 29:1-60.

  • 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.

  • Ayala-Berdon, J., R. Galicia, C. Flores-Ortíz, R. A. Medellín, and J. E. Schondube. 2013. Digestive capacities allow the Mexican long-nosed bat (Leptonycteris nivalis) to live in cold environments. Comparative Biochemistry and Physiology Part A 164:622-628.

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  • Baker, R. J., J. K. Jones, Jr., and D. C. Carter, editors. 1976. Biology of bats of the New World family Phyllostomatidae. Part I. Spec. Publ. Mus. Texas Tech Univ. (10):1-218.

  • Baker, R. J., J. K. Jones, Jr., and D. C. Carter, editors. 1977. Biology of bats of the New World family Phyllostomatidae. Part II. Spec. Publ. Mus. Texas Tech Univ. (13):1-364.

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  • 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> (Accessed April 1, 2015)

  • Brown, C. M. 2008. Natural history and population genetics of the endangered Mexican long-nosed bat, Leptonycteris nivalis (Chiroptera: Phyllostomidae). M. S. thesis, Angelo State University, San Angelo, Texas.

  • Campbell, L. 1995. Endangered and Threatened Animals of Texas: Their Life History and Management. Texas Parks and Wildlife Department, Endangered Resources Branch, Austin, Texas. ix + 129 pp.

  • Cockrum, E. L., and Y. Petryszyn. 1991. The long-nosed bat, Leptonycteris: an endangered species in the southwest? Occas. Pap. Mus. Texas Tech Univ. (142):1-32.

  • Davis, W. B., and D. C. Carter. 1962. Review of the genus Leptonycteris (Mammalia: Chiroptera). Proceedings Biological Society Washington 75:193-197.

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  • Gardner, A. L. 1977c. Feeding habits. Pages 293-350 in R. J. Baker, et al., eds. Biology of bats of the New World family Phyllostomidae. Part II. Special Publication. Museum Texas Tech Univ. (13):1-364.

  • Hensley, A. P., and K. T. Wilkins. 1988. Leptonycteris nivalis. Mammalian Species 307:1-4.

  • Hoyt, R. A., J. S. Altenbach, and D. J. Hafner. 1994. Observations on long-nosed bats (Leptonycteris) in New Mexico. Southwestern Naturalist 39:175-179.

  • Jones, J. K., Jr., and D. C. Carter. 1976. Annotated checklist, with keys to subfamilies and genera. Pages 7-38 in R. J. Baker, J. K. Jones, Jr., and D. C. Carter, editors. Biology of bats of the New World family Phyllostomatidae. Part I. Spec. Publ. Mus. Texas Tech Univ. (10):1-218.

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