6227_Therya

Diversity patterns (henceforth, DPs) exist in all ecosystems around the planet and are constantly changing due to the interaction of abiotic and biotic factors in ecosystems (Chesson 2000; Dirzo and Raven 2003; Brown et al. 2004; Sibly et al. 2012; Villalobos and Rangel 2014). These changes are generally reflected in the fluctuations in abundance, diversity, or richness of the species distributed in a given area. A clear example occurs in urban areas (i.e. geographic spaces with human activity and presence, sensu Weeks) (Weeks 2010), where these types of environments may alter the habitat and therefore, the species composition and dynamics (Rosenzweig 1995; Challenger and Dirzo 2009; Faeth et al. 2011).

Bats are cosmopolitan, they are vagile and present different functional traits, which allows them to be distribute in different ecosystems, including urban environments where they constitute a key component of the mammalian fauna (Van der Ree and McCarthy 2005). It has been observed that in urban environments richness decreases for most bat species, whereas abundance increase only for some groups that are able to adapt to the new characteristics of the environment (generalist species) (Segura et al. 2007; Jung and Kalko 2011; Clavel et al. 2011; Threlfall et al. 2012; Büchi and Vuilleumier 2014; Jung and Threlfall 2016). However, within the urban matrix, suburban areas (i.e. areas of lower human population density located on the periphery of cities or urban areas) have shown a higher bat species richness and abundance compared to urban areas sensu Weeks (2010).

The presence of bats could be related to the availability of food, shelter and foraging sites typical of urban environments (Violle et al. 2007). Furthermore, the modification of these sites and their characteristics can alter bat diversity and consequently generate certain diversity patterns (Russo and Ancillotto 2015). In general, suburban areas have been shown to support higher bat diversity and activity due to increase of potential prey’s number, for generalist and specialist species equally (Shochat et al. 2004; Coleman and Barclay 2012; Luck et al. 2013).

Three hypotheses may explain this phenomenon: i) the heterogeneity hypothesis, ii) the intermediate disturbance hypothesis, and iii) the habitat productivity hypothesis. Together they describe how richness and abundance vary with the disturbance frequency and intensity. Disturbance creates heterogeneity in the environment that, combined with the addition of anthropogenic organic matter, increases primary productivity and provides a greater number of available resources (Connell 1978; Shochat et al. 2004; Shochat et al. 2006; McKinney 2008; Gaston and Gaston 2010; Threlfall et al. 2011). Such DPs have been observed in some vertebrate groups, such as bats (Duchamp et al. 2004).

In Mexico most studies aimed at characterizing bat diversity in urban or suburban environments, have focused on tropical regions (Medellín 1993; Arita 1993), even though, more than 50% of the national territory has a dry or semi-dry climate type, where such studies are scare (SEMARNAT 2015). The city of Zacatecas is characterized by semi-dry climate, and the only available information about bat diversity comes from the company URSAMEX (2014), which was the responsible for the construction of the suburban park “Ecoparque Centenario” (ECO), our study area. They report 5 bat species: Mormoops megalophylla, Leptonycteris nivalis, Myotis auriculus, M. planiceps and Dermanura azteca, although 3 species distribution (L. nivalis, M. planiceps and D. azteca) does not correspond to what has been previously reported (Medellín et al. 2008; Ortega et al. 2022). In addition, neither the identification methods nor the sampling effort is presented, so the information is incomplete and inaccurate. The ECO is surrounded by active mines, and this site was recognized as a natural protected area (URSAMEX 2014); it has been suggested that mines can be used by different species of bats as perching sites, which could favor their diversity. On the other hand, in addition to the recognition, the area requires constant and exhaustive diversity studies. Therefore, our aims were to determine the bat diversity using 2 identification methods (acoustic and morphologic) and to estimate diversity index in order to establish the patterns present in a suburban area “Ecoparque Centenario” belonging to the Mexican plateau during an annual cycle.

Materials and Methods

The Mexican plateau is located between the Western and Eastern Sierras, and, in the south, it is limited by the transversal volcan axis. This is an extensive area characterized by altitudes near 2000 m asl. The predominant type of vegetation is xeric scrub, pine-oak forest and isolated patches of low deciduous forest (Rzedowski 2006). The ECO, protected natural area belonging to the Mexican plateau, is located in the Arroyo de la Plata micro-watershed, between the Central Mesa and Western Sierra Madre physiographic regions. The ECO is located at coordinates: 22º 46’ 49.14” N, 102º 32’ 37.96” W (Figure 1), between Zacatecas, Vetagrande and Guadalupe municipalities; the park is in the border area of the Zacatecas city, at an altitude of 2448 m asl, with an average annual rainfall of 400 mm to 450 mm. The climate type corresponds to BS1kw (dry or semi-dry with temperate regions and an average annual temperature that ranges between 12 and 18 ºC; García 2004).

The predominant vegetation is induced grassland, riparian vegetation, xerophytic scrub and Opuntia spp. scrub (Rzedowski 2006). The tree density is composed by pirul (Schinus molle) and mesquite (Prosopis sp.) along the stream banks (URSAMEX 2014). Six sampling points were selected near to water bodies and alternated according to the annual season to increase the probability of bat capture during the dry season. Four mist nets (2.5 × 6 m) were placed alternately at the 6 points (Bell 1980; Kurta and Kunz 1988; MacSwiney et al. 2008; Gilley and Kennedy 2010). Mist nets were sampled for 5 days, each month, for one year, from April 2022 to May 2023. They were opened during 5 hours, after sunset with monitoring every 30 minutes (Holloway and Barclay 2000; MacSwiney et al. 2008; Coleman and Barclay 2012; Barboza-Marquez et al. 2014).

For acoustic monitoring, the Echo Meter Touch 2 Pro ultrasonic microphone (connected to a tablet Lenovo Xiaoxin Pad 2022) and Echo Meter software (Wildlife Acoustics, Inc. Maynard, Massachusetts) were used to record EPs. The detection range of echolocation calls was set to a minimum frequency of 15,000 Hz with a sampling rate of 384 kHz (Pettersson 2004). Acoustic monitoring was semi-active and was conducted by walking the trail from net to net and lasted as long as the mist nets remained open (five hours per day; MacSwiney et al. 2020).

BatSound V4.1 software (Pettersson 2004) was used to characterize search phases of the EPs, as they are relatively constant compared to other types of bat vocalizations (e.g. social pulses, feeding buzzes) (Fenton and Bell 1981; O’Farrell and Miller 1999; Barclay 1999; Papadopoulos and Allen 2007; Agranat 2013). EPs with an intensity less than 30 dB were not considered for characterization, since it has been determined that frequencies less than this value tend to attenuate at short distances, therefore higher intensity frequencies can travel farther in the environment and consequently, be recorded by ultrasonic microphones (Surlykke and Kalko 2008). These parameters (measured in kHz) were: maximum frequency (Fmax), minimum frequency (Fmin), peak frequency, and bandwidth (the difference between Fmax and Fmin), whereas intensity was expressed in dB, and duration (DUR) in milliseconds (ms) (Corben 2004; Miller 2004). The values of each pulse were checked against the “Compendio de Llamados de Ecolocalización de los murciélagos insectívoros mexicanos” (Ortega et al. 2022) and the SONOZOTZ echolocation call library (Zamora-Gutiérrez et al. 2020). A species was assigned under the concept of sonospecies in the case of meeting the above assumptions, particularly Fmin, peak frequency and DUR (Thomas et al. 1987). Morphological bat identifications were made using the field keys in Medellín et al. (2008), according to the diagnostic morphological characteristics (Martínez-Rodríguez et al. 2024).

Presence/absence data matrices were constructed for the species identified by both methods. The diversity estimators ICE (Incidence-based Coverage Estimators) and Chao 2 (Lee and Chao 1994) were calculated using the EstimateS v 9.1 software (Colwell 2013). Species accumulation curves were generated using the PAST v 4.17 software (Hammer and Harper 2001) with i) acoustic and morphological data, ii) the total observed data for the number of species and 95% confidence intervals, and iii) the non-parametric estimators (ICE and Chao 2), acoustic, morphological and the total data observations with both methods (Chao et al. 2009). Shannon, Margalef, Gini-Simpson and Berger-Parker indices were calculated (Margalef 1972; Moreno 2001; Magurran 2007; Magurran et al. 2019) in order to elucidate the diversity patterns in the suburban area.

Results

The total sampling effort was 59 days, 17,700 net-hours and 10,215 acoustic recordings of which 3,068 met the characteristics described in the methodology. Twenty-eight species, 14 genera and 4 families were identified (Table 1). Only 19 species were identified by analysis of their EPs (Table 2); 8 species were identified using both methods and one species was identified only by taxonomic keys (Choeronycteris mexicana; Figure 2). The family Vespertilionidae was the most represented with 9 genera and 20 species (71.4 %), with 7 species correspond to the genus Myotis (25 %) and 5 to the genus Lasiurus (17.8 %). Three genera and 6 species were included in the Molossidae family (21 %), and the most represented genus was Nyctinomops with 4 species (14.2 %). Only one species was recorded in the families Mormoopidae and Phyllostomidae, Mormoops megalophylla and C. mexicana, respectively (7.1 %).

In the context of acoustic monitoring, the species with highest number of occurrences were Lasiurus intermedius (n = 630), Myotis yumanensis (n = 341), Tadarida brasiliensis (n = 341), L. cinereus (n = 330), M. volans (n = 262), M. californicus (n = 237), and N. laticaudatus (n = 200). The species with the fewest recorded occurrences were Corynorhinus mexicanus (n = 1), Rhogeessa parvula (n = 2), and M. megalophylla (n = 2). We also recorded L. ega (Figure 2 B, e), which has not been previously registered in the north-central region of the country.

For the morphological analysis, 37 specimens were captured which corresponded to 3 families, 5 genera and 9 species; 35 specimens and 7 species belong to the Vespertilionidae (94.5%): M. yumanensis (n=10), M. californicus (n=11), M. ciliolabrum (n=3), M. volans (n=3), L. ega (n=1), L. frantzii (n=1), and C. townsendii (n=6). In the families Molossidae and Phyllostomidae only 1 species was captured: T. brasiliensis (n=1) and C. mexicana (n=1), respectively. The photographs and EPs corresponding to each species are shown in Figure 2, except for C. mexicana (Figure 2 B, i) considered a “whispering” species due to its EP’s characteristics (low intensity and high frequency).

The first accumulation curve (Figure 3 A) shows the differences in the number of species recorded between the identification methods used. While acoustic monitoring (blue line = 27) tends to asymptote, morphological identification (red line = 9) shows no signs of saturation. Figure 3 B shows the total data (green line = 28), which indicates that both methods cover 87.96% of diversity according to the upper confidence interval (blue line = 31.83). However, figure 3 C shows that the sampling effort was satisfactory according to the diversity estimators ICE (blue line = 30.12) and Chao 2 (pink line = 29.47), which suggest that between 92.89 % and 95 % of the richness was recorded in the study area. Acoustic monitoring (cyan line = 27) recovered between 89.58% (ICE) and 91.6% (Chao 2) of the richness. Morphological identification (orange line = 9) recorded between 29.86% (ICE) and 30.5% (Chao 2) of the richness.

The value for Margalef specific diversity index was R = 4.421, which indicates a moderate diversity. The Shannon diversity index had a value of H’ = 3.073, which also indicates moderate diversity in the study area. For the Gini-Simpson index, a value of 1-D = 0.9474 was obtained; this value indicates that there is a high probability of obtaining two different species from a random sample, thus indicating that there is no species dominance in the study area.

Finally, the value obtained for the Berger-Parker index was D = 0.09131, indicating that the species with the highest proportion of occurrences in the sample represents 9.1% of the recorded richness, therefore, there is no indication of dominance of any species in the study area. With these values, together with the bat diversity composition, we can infer that the ECO complies with the hypothesis of intermediate disturbance.

Discussion

The most represented bat family in our sample was Vespertilionidae (20 spp.), followed by the Molossidae (6 spp.). The diversity composition recorded in this study is consistent with previously described diversity patterns in arid and semiarid climates of the Mexican plateau and the south of Arizona (USA) (e.g. Ortega and Arita 1998; López-González et al. 2015; Segura-Trujillo et al. 2016; Bazelman 2016; Dwyer 2021; Segura-Trujillo et al. 2022; Ramos-H et al. 2024). Because the study site is nearby to the transitional zone between the Nearctic and Neotropical biogeographic regions, it is possible to find elements of Neotropical origin, such as species of the families Molossidae, Mormoopidae, and Phyllostomidae (Ortega and Arita 1998; López-González et al. 2015). In this study these elements represent only 8 species (i.e. 28.5% of the total sample).

In relation with the generalist and specialist bat species, previous studies have determined that at least 10 of the 28 identified species in this study are generalists, whose presence is correlated with suburban environments (T. brasiliensis, E. fuscus, M. yumanensis, M. californicus, M. velifer, M. volans, N. macrotis, L. xanthinus, L. intermedius and M. megalophylla) (Avila-Flores and Fenton 2005; Dixon 2012; Bazelman 2016; Rodríguez-Aguilar et al. 2017; Adams 2021; Dwyer 2021; Dwyer et al. 2024; Briones-Salas et al. 2024). However, it has been reported that the species N. femorosaccus, C. townsendii, N. brasiliensis, M. auriculus, P. hesperus and C. mexicana have specialized habits, with relatively low activity levels in urban areas (Husar 1976; Arroyo-Cabrales et al. 1987; Bazelman 2016; Rodríguez-Aguilar et al. 2017; Dwyer 2021; Dwyer et al. 2024). In addition to having specialist habits, C. mexicana has been classified as near threatened by the IUCN (Solari 2018). Therefore, this work contributes to generating information to make decisions about conservation strategies for species that can inhabit suburban habitats in the Mexican plateau.

Evidence suggests that species of the genus Lasiurus (L. frantzii, L. ega, L. cinereus and L. intermedius) tolerate intermediate levels of urbanization. However, given their foraging and refuge site characteristics, they tend to avoid such environments. The exception is L. xanthinus, which has not been reported reduce its presence in habitats due to increased urbanization (Aguilar et al. 2013; Dwyer 2021).

According to Moreno and Halffter (2001), it is necessary to recover a minimum of 90 to 95 % of the bat diversity to ascertain that the sampling effort was sufficient. We recovered between 92.89 % (ICE) and 95 % (Chao 2), indicating a satisfactory sampling effort. This is due to the use of two identification methods, which have been deemed optimal in suburban environments or areas characterized by minimal vegetation cover, such as xerophytic scrub vegetation (Rautenbach et al. 1996; Kuenzi and Morrison 1998; Rydell et al. 2002; Berry et al. 2004; MacSwiney et al. 2020). Furthermore, these methods are complementary to each other, and their efficiency varies depending on habitat characteristics and the trophic guild to which the bat species belong (i.e. open space aerial foragers, closed space aerial foragers, surface foragers, and edge space foragers). For instance, numerous authors (e.g. O’Farrell and Miller 1999; Kalko et al. 2008; MacSwiney et al. 2008) have mentioned that acoustic monitoring has been found to be the most efficient in recording species that forage in open spaces, while mist nets have been determined to be optimal for capturing species that forage in closed or surface spaces (La Val 1970; Kunz 1973; Kunz and Brock 1975; Kuenzi and Morrison 1998; Rydell et al. 2002; Larsen et al. 2007). This distinction is evident in the accumulation curves of each method and the feeding habits reported in previous studies (Mora-Villa et al. 2014; Segura-Trujillo et al. 2016). Acoustic monitoring recorded 13 genera and 27 species (96.4% of the sample), of which six molossid species belonged to the guild of open-space aerial foragers and at least 17 species of Vespertilionids belonged to the guild of edge-space foragers.

In the other hand, the morphological identification recorded 9 spp. of which 8 were also identified through their EPs, and which mostly belong to the guild of closed-space foragers and edge-space foragers. The only exception was C. mexicana which feeds on pollen and nectar from Agavaceae flowers and has EPs that are complicated to record, like other Phyllostomids species, but they are relatively easy to capture with mist nets (Kunz and Kurta 1988; Simmons and Voss 1998; Clarke et al. 2005; MacSwiney et al. 2008; Pérez-Hernández and Martínez-Coronel 2023). In addition, the combined methods allowed us to identify acoustically and morphologically the species L. ega, which some authors have reported that it is distributed mainly on the slope of the Gulf of Mexico (Medellín et al. 2008; Barquez and Diaz 2016; Ortega et al. 2022), although others mention that its distribution includes the north-central region of the country (Kurta and Lehr 1995). Because of this uncertainty and the migratory habits of this species, it is imperative to provide information on the distribution of the species in the arid and semiarid regions of the country, where studies are limited.

In the context of acoustic characterization, it was observed that the Fmin values recorded for all the 27 species matched the values of the “Compendio de Llamados de Ecolocalización de los murciélagos insectívoros mexicanos” and the SONOZOTZ echolocation call library (Ortega et al. 2022). The recorded Fmax values differ in 5 species (Lasiurus xanthinus, L. cinereus, M. megalophylla, Molossus nigricans and N. laticaudatus). Conversely, peak frequency values differ in one species (Corynorhinus townsendii) and the recorded DUR values differ in 17 spp. (Antrozous pallidus, Baeodon alleni, C. mexicanus, C. townsendii, Eptesicus fuscus, L. cinereus, L. ega, L. frantzii, M. ciliolabrum, M. auriculus, Parastrellus hesperus, R. parvula, N. aurispinosus, N. femorosaccus, N. laticaudatus, N. macrotis and T. brasiliensis).

The parameters that exhibited the most discrepancy was Fmax and DUR, which may be associated with the capacity of bats to decrease Fmax and increase DUR in response to the climatic and structural characteristics of the site. This phenomenon can be a strategy employed by bats to mitigate atmospheric attenuation and avert the masking of their EPs by anthropogenic sounds in this context (Thomas et al. 1987; Wund 2006; Gillam et al. 2009).

The calculated diversity indices values suggest that the study area has a moderate bat diversity (H’ = 3.073, R = 4.421, 1-D = 0.9474 and, D = 0.09131). There is a possibility that this is due to the suburban park characteristics (i.e. moderate levels of urbanization, tree cover, water bodies and streetlights) and the presence of caves and abandoned mines in the vicinity of the ECO. Several studies have documented that these features may explain why the richness of bat species was higher in suburban parks or in suburban areas due the presence of available roost and foraging sites for different species of bats (Kurta and Teramino 1992; Gehrt and Chelsvig 2008; Loeb et al. 2009; Russo and Ancillotto 2015). In addition, we inferred from the calculated values of each index and the suburban characteristics of the environment, the diversity pattern present in the ECO polygon corresponds to the hypothesis of intermediate disturbance.

This hypothesis states that species can take advantage of moderately altered habitats and increase the richness and diversity in general (Connell 1978; Castro-Luna et al. 2007; Threlfall et al. 2011; Dodd et al. 2012). Our study is an example of how systematic and formal studies about diversity of bats using 2 methods of identification, can elucidate and provide information about the species distribution and their EPs characteristics in regions where these types of studies are limited. In addition, it is important to highlight the importance of its implementation to reduce bias regarding the diversity of bats present. In our case, for example, the company URSAMEX (2014) mentions that they identified five species of bats, but three species do not correspond to the reported distribution, moreover statistical methods were not used to evaluate the sampling effort, and only one identification method was used, which could have underestimated the diversity of bats present in the park. In fact, we only identified two of these mentioned species which correspond to Myotis auriculus and Mormoops megalophylla. This is an example of how the use of complementary methods for bat species identification can expand and provide accurate information about the actual knowledge of diversity at the local level in this type of environment which cover a Mexican plateau.

Conclusions

We registered 28 bat species with two methods of identification. Our study represents the first formal and systematic listing of bat species in a suburban environment in the Mexican plateau region and particularly in the state of Zacatecas. In addition, we registered the species L. ega which was not reported in the region. Finally, our sampling effort was satisfactory, and the bat diversity pattern identified in the ECO corresponds to the pattern observed in the north-central region of the country, that is, a greater representation of the families Vespertilionidae and Molossidae. Furthermore, according to the calculated diversity indexes values, it is inferred that the suburban characteristics of Park maintain a moderate diversity of chiropteran species and it is suggested that it corresponds to the pattern of the intermediate disturbance hypothesis.

Acknowledgments

We thank the reviewers of this work whose invaluable comments greatly improved the quality of this work. We also thank COZCyT who provided us with financial support to present our study in different forums, as well as the Secretaría Agua y Medio Ambiente (SAMA) for allowing us to carry out the study in the “Ecoparque Centenario”.

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Associated editor: Sergio Solari

Submitted: August 21, 2025; Reviewed: November 15, 2025

Accepted: December 2, 2025; Published on line: December 17, 2025

THERYA, 2026, Vol. 17(1):45-58

DOI:10.12933/therya-25-6227 ISSN 2007-3364

Figure 1. Geographic location of the “Ecoparque Centenario” (ECO) area. The Mexican Plateau is shown as the red silhouette. The state of Zacatecas is shown as black outline. The municipality of Zacatecas is shown as bold black line (b) and yellow line (c-d). The Ecoparque Centenario is shown as red dot (b-c) and white line (d)

Figure 2. Echolocation calls (A) and photographs (B) of bat species captured. a) M. yumanensis, b) M. californicus, c) M. volans, d) M. ciliolabrum, e) L. ega, f) L. frantzii, g) C. townsendii, h) T. brasiliensis, and i) C. mexicana.

Table 1. List of bat species identified.

Family	Genus	Specie	Morphologic presences	Individuals captured	Acoustic presences	Acoustic records	Total presences	Total detections
Vespertilionidae	Antrozous	pallidus	0	0	4	6	4	6
	Baeodon	alleni	0	0	8	21	8	21
	Corynorhinus	mexicanus	0	0	1	1	1	1
		townsendii	4	6	8	23	11	29
	Eptesicus	fuscus	0	0	16	94	16	94
	Lasiurus	cinereus	0	0	32	330	32	330
		ega	1	1	8	48	9	49
		frantzii	1	1	14	36	14	37
		intermedius	0	0	41	623	41	623
		xanthinus	0	0	3	8	3	8
	Myotis	auriculus	0	0	22	127	22	127
		californicus	8	11	32	237	34	248
		fortidens	0	0	18	79	18	79
		ciliolabrum	3	3	19	53	20	56
		velifer	0	0	19	67	19	67
		volans	3	3	28	262	28	265
		yumanensis	6	10	31	341	35	351
	Neoeptesicus	brasiliensis	0	0	12	52	12	52
	Parastrellus	hesperus	0	0	4	9	4	9
	Rhogeessa	parvula	0	0	2	2	2	2
Mormoopidae	Mormoops	megalophylla	0	0	1	2	1	2
Molossidae	Molossus	nigricans	0	0	13	26	13	26
	Nyctinomops	aurispinosus	0	0	20	43	20	43
		femorosaccus	0	0	8	13	8	13
		laticaudatus	0	0	30	200	30	200
		macrotis	0	0	11	24	11	24
	Tadarida	brasiliensis	1	1	32	341	32	342
Phyllostomidae	Choeronycteris	mexicana	1	1	0	0	0	1
Total	14	٢٨	28	37	437	3068	449	3105

Table 2. Characterization of echolocation pulses (EPs) from bat species identified.

Family	Genus	Specie	Maxium frequency (kHz)	Minium frequency (kHz)	Peak frequency (KHz)	Duration (ms)	Bandwidth (kHz)	n
Vespertilionidae	Antrozous	pallidus	59.1 ± 17.3	27.8 ± 2.8	36.7 ± 6.7	9.3 ± 2.6	31.3 ± 14.6	6
	Baeodon	alleni	98.5 ± 4.4	33.3 ± 2.1	46.5 ± 1.3	5.1 ± 0.6	65.2 ± 5.3	21
	Corynorhinus	mexicanus	46.9 ± 0.0	21.6 ± 0.0	33.3 ± 0.0	12.3 ± 0.0	25.3 ± 0.0	1
		townsendii	40.1 ± 2.4	23.9 ± 1.9	31.2 ± 1.3	11.7 ± 2.9	16.1 ± 2.6	23
	Eptesicus	fuscus	51.4 ± 5.3	27.7 ± 1.9	33.5 ± 1.9	9.8 ± 1.8	23.7 ± 5.8	94
	Lasiurus	cinereus	49.8 ± 6.9	25.6 ± 1.9	32.7 ± 2.6	11.9 ± 2.4	24.2 ± 6.8	330
		ega	55.8 ± 11.7	34 ± 4.8	38.2 ± 5.4	9.7 ± 4.5	21.7 ± 8.2	48
		frantzii	84.6 ± 13.9	37.3 ± 2.4	45.5 ± 2.3	5.9 ± 1.9	47.3 ± 15.3	36
		intermedius	41.2 ± 6.6	23.4 ± 1.5	28.2 ± 1.6	12.7 ± 2.2	17.9 ± 6.7	623
		xanthinus	72.9 ± 7.3	32.8 ± 1.0	39.1 ± 1.2	7.8 ± 0.7	40.1 ± ٧.٩	8
	Myotis	auriculus	86.6 ± 8.9	32.2 ± 2.2	43.9 ± 1.8	6.2 ± 1.0	54.4 ± 9.5	127
		californicus	94.9 ± 5.3	41.8 ± 2.3	54.3 ± 2.3	5.1 ± 2.7	53.1 ± 6.2	237
		fortidens	98.7 ± 4.6	42.9 ± 1.6	55.9 ± 1.2	4.4 ± 0.6	55.7 ± 5.1	79
		ciliolabrum	100.6 ± 5.2	39.7 ± 2.5	54.9 ± 3.6	4.6 ± 0.9	60.8 ± 5.2	53
		velifer	83.5 ± 11.3	37.8 ± 1.7	46.1 ± 2.9	5.7 ± 0.6	45.6 ± 11.1	67
		volans	93.1 ± 6.0	37.5 ± 2.7	47.7 ± 2.7	5.5 ± 0.7	55.6 ± 6.5	262
		yumanensis	96.9 ± 5.6	42.7 ± 2.8	54.6 ± 3.9	4.7 ± 0.6	54.2 ± 6.0	341
	Neoeptesicus	brasiliensis	53.7 ± 4.1	30.8 ± 1.9	37 ± 2.0	10.9 ± 1.7	22.9 ± 4.5	52
	Parastrellus	hesperus	69.9 ± 4.4	41.9 ± 0.9	47.1 ± 1.8	6.2 ± 2.3	27.9 ± 3.7	9
	Rhogeessa	parvula	87.9 ± 0.6	41.7 ± 1.5	53.9 ± 0.0	3.8 ± 0.2	46.2 ± 2.1	2
Mormoopidae	Mormoops	megalophylla	57.6 ± 1.2	41.5 ± 0.7	54 ± 0.2	6.4 ± 1.0	16.1 ± 1.8	2
Molossidae	Molossus	nigricans	35.1 ± 3.7	25.7 ± 1.6	30.1 ± 1.3	13.3 ± 2.3	9.4 ± 3.4	26
	Nyctinomops	aurispinosus	36.2 ± 6.1	19.7 ± 0.8	26.4 ± 1.8	14.3 ± 1.7	16.4 ± 6.3	43
		femorosaccus	32.9 ± 4.7	18 ± 1.0	23.9 ± 0.6	13.2 ± 1.9	14.9 ± 4.7	13
		laticaudatus	35.4 ± 6.6	20.7 ± 1.0	25.6 ± 1.8	13.8 ± 2.2	14.7 ± 6.8	200
		macrotis	27.7 ± 3.3	17.4 ± 2.8	21.5 ± 1.9	14.9 ± 2.3	10.3 ± 4.7	24
	Tadarida	brasiliensis	40.7 ± 8.6	24.1 ± 2.3	29.5 ± 3.7	12.8 ± 2.6	16.5 ± 7.3	341
Total								3068