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Study of Roost Selection and Habits of a Bat Hipposideros armiger in Mainland China.

Byline: Yanzhen Bu Meixin Wang Chan Zhang Haixia Zhang Lezhen Zhao Huixian Zhou Yan Yu and Hongxing Niu

Abstract

Roost selection perching activity habits and the degree of human disturbance of a bat Hipposideros armiger were studied from December 2010 to September 2013 in mainland China. Sixty-nine potential roosts were investigated. Among those 54 roosts were occupied by bats of which 20 roosts were used by H. armiger. The differences of various roosts were compared by measuring the structural characteristics of these roosts the microenvironment and the degree of human disturbance. Comparing the 54 roosts used by bats with the 15 roosts not used by bats it was found that the former had fewer entrances larger volumes longer cave lengths higher temperatures and lower illuminance. Fifteen hibernation roosts of H. armiger had relatively lower illuminance and fewer entrances. Eleven breeding roosts of H. armiger had relatively longer cave lengths and lower illuminance whereas the temperature and humidity were relatively high. H. armiger do not often co-inhabit roosts with other species.

However sometimes they co-inhabit roosts with Hipposideros pratti. Pregnant females give birth to a single young each year between May and early June. During the breeding and young rearing periods the males and females usually live in different roosts except for a few males that remain with the females in the breeding place.

Key words: Roost selection Hipposideros armiger conservation degree of human disturbance.

INTRODUCTION

Bats spend over half of their lives in roosts which provide them with protection and sites for resting mating hibernation rearing young and social interactions (Kunz 1982). Although many bat species use caves only as an alternate refuge some species rely completely on caves for day roosting. In Mexico for example 45% (60 of 134) of bat species are cave dwellers with 27 using caves as the main roost and 33 additional species using caves occasionally (Arita 1993). In Europe caves are used regularly or occasionally by 46 bat species (Nagy and Postawa 2011).

Roost selection by bats depends on many factors including temperature humidity air flow light intensity safety from predators proximity to foraging areas and take-off height (Morrison 1980; Tuttle and Stevenson 1981; Kunz 1982; Hill and Smith 1984; McCracken 1989). Occupation of roost sites with an appropriate microclimate can minimize the energetic costs related to thermoregulation food digestion and assimilation maintenance of a permanent state of alertness (which allow bats to avoid predation and to interact socially) gestation embryonic development parental care lactation and spermatogenesis (Kunz 1973; Humphrey 1975; Tuttle and Stevenson 1981; McNab 1982; Hill and Smith 1984; Bonaccorso et al. 1992; Hamilton and Barclay 1994).

The microclimate of a cave is dependent on latitude longitude and altitude as well as the length of the caves the number of entrances and the average temperature (Kowalski and Archeologiczne 1954). The number of bat species using a cave is correlated with the length of the cave (Arita 1996) the density of underground sites in each area (Brunet and Medellin 2001) and environmental factors such as geographical location and temperature range (Rehak 2006; Ulrich et al. 2007).

Most bats use a variety of roosts including man-made structures (Kunz 1982; Fenton 2001). Many species can endure a wide range of roost conditions. The roosting ecology of bats has been well studied for species in the temperate zone. Bats may use different roosts according to different requirements for environmental conditions in different seasons. In the winter most temperate zone bats hibernate in cooler roosts so they can survive through the period of cold and food shortages (Kurta 1986). In the summer or the breeding season maternity colonies usually exist in roosts with higher ambient temperatures (Henshaw 1960; Betts 1997; Entwistle et al. 1997; Williams and Brittingham 1997). Reproductive females which need to maintain a higher body temperature to facilitate fetal growth may take advantage of higher ambient temperatures in roosts to reduce metabolic energy expenditure (McNab 1982).

In contrast males or non-breeding females which do not have the pressure of maintaining a higher body temperature for fetal growth choose roosting sites with lower ambient temperatures and frequently use torpor to reduce metabolic energy expenditure (Hamilton and Barclay 1994).

Bats are sensitive to climate change and roost deterioration and have been recognized as valuable bio-indicators (Jones et al. 2009). The suitability and availability of roosts may influence the survival reproduction and distribution of bats (Humphrey 1975). Because of cave exploitation for tourism the extensive use of pesticides the demolition of many old buildings and the inclusion of bats in the diet bat populations in China appear to have decreased considerably in the last 30 years (Zhang et al. 2009).

The great leaf-nosed bat Hipposideros armiger (Hipposideridae Chiroptera) one of the largest species within the genus Hipposideros typically roosts in caves and feeds in open spaces in woodlands gardens and around trees (Bates and Harrison 1997). It is characterized by high wing loading low aspect ratio and average wing tip shape index (Wei et al. 2011). The wing loading is a measure of the surface area of the wings compared to the body weight. The aspect ratio describes the shape of the wings and a low aspect ratio corresponds with shorter wings and less efficient flight. The wing tip shape index quantifies the pointedness of the wing tips (Jennings et al. 2004). The flight behavior of bats is correlation with wing loading aspect ratio and wing tip shape index (Norberg and Rayner 1987). H. armiger has a wide Asian geographical distribution that includes India Nepal Myanmar Vietnam Laos Cambodia Thailand China and the Malay Peninsula (Simmons et al. 2005; Bates et al. 2007).

In mainland China it is widely distributed in 13 provinces of South China (Wang 2003). Studies on H. armiger have been conducted on the relationships between eco-morphology and prey selection (wang et al. 2005) echolocation calls (Bogdanowicz et al. 1999; Zhao et al. 2003) karyology analysis (Gu 2001; Wu et al. 2003) some mitochondrial DNA sequences (Li et al. 2006) characterization of microsatellite loci (Guo et al. 2008) phylogeography (Lin et al. 2013) evolutionary analysis (Chen et al. 2014) and roost selection in Taiwan (Ho and Lee 2003). However in mainland China the conservation state of their roosts the habits of the species the role of cave microclimate and other environmental factors influencing the distribution of this species are poorly known.

South China is characterized by numerous high mountains deep valleys large rivers and environmental heterogeneity and harbors many species (Li and Fang 1999). To understand the microclimate and structure of caves the surrounding habitat disturbance by humans and relative abundance of H. armiger and to examine the differences in the environmental conditions of caves used by the species in different seasons 69 underground sites in nine provinces in south China were investigated in a 3-year study (from December 2010 to September 2013). Such information is critical for improving the conservation management of this species and for determining whether economic development and biological conservation are indeed compatible.

MATERIALS AND METHODS

Study sites

This research was conducted in nine provinces in south China (N1830'21"-3026'12" E9849'31"-11815'10"). Among these provinces Hainan is located in a tropical zone. Jiangxi Fujian Hunan Guizhou and Sichuan Provinces are situated in a subtropical zone. Yunnan Guangdong and Guangxi Provinces stretch across tropical and subtropical zones (Fig. 1).

Roost characteristics

Sixty-nine potential roosts were found using previous records and field survey. Fifty-four of these roosts are currently being used by various species of bats including H. armiger. All of the roosts in natural caves and other man-made underground structures such as abandoned mines and tunnels are located in a mountainous area and are nearby natural forests or secondary forests. All fieldwork abided by the Law of the People's Republic of China on the Protection of Wildlife. The fieldwork was conducted during four periods: December 2010March 2011 JuneSeptember 2011 December 2011March 2012 and JulySeptember 2013. Each cave was visited at least twice (once in the summer period when the bats are active and once during their hibernation period).

Nineteen structural variables were recorded for each of the 69 potential roosts. These include the elevation of the roost the maximal height and width of the passage the number of entrances and the number of chambers the orientation of the entrances the height and width of the entrances the total lengths of all tunnels the average height and width of all tunnels the volume of the roost the distance to the nearest water source the total floor area covered by water inside the roost the air temperature and relative humidity in the tunnels the longitude and latitude of the entrances and the light intensity.

At each visit we measured the air temperature and relative humidity using a digital thermo-hygrometer (Guangdong Benetech GM1360; precision 1% and relative humidity: 3C). Light intensity was measured with a light meter (precision: 0.1 lux Hong Kong Smart sensor AR823) at sites where bats were present or at equivalent sites within roosts that were not being used by bats. The light intensity was divided into three levels: (1) greater than 10 lux (2) 0.1 lux10 lux and (3) less than 0.1 lux. The area covered by water was classified into four levels: (1) greater than 85% (2) 50%85% (3) 15%50% and (4) less than 15%.

For the caves that are used for tourism the frequency of human activities in the caves and the degree of disturbance to the underground roosts are categorized into three levels: (1) serious disturbance (exploited for tourism burning incense fireworks and firecrackers) (2) light disturbance (burning incense and occasional visiting by people) and (3) no disturbance (no human activities in the caves). We also measured the distance of each roost to both the nearest road and the nearest building as an index of disturbance.

Roosts used by H. armiger were categorized into hibernacula and summer roosts. Summer roosts were further divided into breeding roosts consisting mainly of reproductive females and non-breeding roosts consisting mainly of males.

Biological data

Bats were captured nightly and daily at entrances or inside caves by using mist nets and hand nets. Calipers and a digital electronic scale (Guangdong Weiheng WH-DS01 accuracy = 0.01 g) were used to take measurements of the length of the forearm the body size the body mass the length of some fingers and the length of the hind foot. We obtained information about sex age (juvenile or adult) and reproductive status (only for females: inactive pregnant or lactating) partly to determine roost use: hibernacula maternity roost or non- breeding male and/or female roosts. Bat species were identified based on their morphological characteristics following the method of Smith et al. (2009). All captured bats were released as soon as possible after being measured.

For measuring the population sizes of H. armiger and other bat species the number of bats in a colony less than 50 individuals were directly counted. For larger clusters and mixed species groups the colony was photographed using a digital camera and the bats were counted from the photos.

Statistical analysis

Data analysis was performed in SPSS 13.0 for Windows. The independent- samples T test was used to determine significant differences in the characteristics of the different roosts. In addition the characteristic data of the roosts were also processed using the principal component analysis method. The Chi-square test was used to determine significant difference in the orientation of the entrances and in the selection of the cave types. Moreover the Chi-square test was also used to evaluate the degree of human disturbance on the roosts.

RESULTS

Roost characteristics and microclimate

Among the 69 roosts 54 were occupied by bats and H. armiger occupied 20 of the roosts. Among these 20 roosts 11 roosts were breeding roosts and nine roosts were non-breeding roosts. Moreover ten roosts had more than 200 H. armiger seven roosts contained between 100 and 200 bats and three roosts contained less than 100 bats. All 20 of these roosts were summer roosts and 15 of them were also used by H. armiger in the winter. Eighteen of the 20 roosts were also occupied by other bat species including Hipposideros pratti Hipposideros larvatus Hipposideros pomona Aselliscus stoliczkanus Rhinolophus rex Rhinolophus ferrumequintum Rhinolophus cornutus Rhinolophus thomasi Rhinolophus pusillus Rhinolophus affinis Myotis ricketti Myotis frater Myotis chinensis Myotis altarium Miniopterus schreibersii Nyctalus noctula Nyctalus velutinus Ia io Pipistrellus abramus and Taphozous melanopogon.

All of the 69 roosts were located at elevations between 41 m and 2098 m. The elevations of the 54 roosts occupied by bats were between 41 and 2098 m. Fifteen hibernacula of H. armiger were between 89 m and 2042 m. Five roosts used by H. armiger only in the summer had elevations between 201 m and 1130 m. Eleven breeding roosts had elevations from 109 to 2042 m and nine non-breeding roosts were between 89 m and 1130 m. The data indicate that the roost selection of H. armiger is unrelated to altitude (Table ). Forty-seven potential roosts had only one entrance 13 had two entrances four had three entrances and five had four entrances. The distances from 35 of these roosts to the nearest permanent water source were within 100 m. Thirty- one roost-water distances were between 100 m and 500 m and three were greater than 500 m (Table ).

Compared the 54 roosts used by bats with 15 roosts without bats it was found that the former had fewer entrances larger volumes longer cave lengths higher temperatures and lower illuminance. When the 20 roosts occupied by H. armiger were compared with the other 34 roosts without this species no significant difference was detected in the physical structure of the roosts light intensity relative humidity and roost temperature (p greater than 0.05). When the 15 hibernation roosts of H. armiger were compared with non-hibernacula roosts there was a significant difference in light intensity (pless than 0.05) and

Table .- Characteristics of the different types of bat roosts investigated in mainland China. Variables are expressed as the means standard deviation (SD).

Variables###Roost type

###HR###OR###HRW###HRS###HB###HNB

Sample size###20###34###15###5###11###9

Elevation of the roost (m)###754.55###928.06###850.53###466.60###1058.09###392.25

###614.45###730.0###656.94###380.40###648.35###318.99

Number of entrances###1.75###1.62###1.27###2.40###1.91###1.56

###1.11###0.88###0.45###1.14###1.22###1.01

Distance to nearest permanent water (m)###131.00###251.47###101.33###220.00###120.00###144.44

###130.25###306.71###105.75###168.07###117.47###150.59

Height of entrances (m)###4.03###9.01###3.64###5.22###4.02###4.04

###2.35###17.87###1.53###3.96###1.55###3.18

Width of entrances (m)###4.53###6.69###4.31###5.20###4.91###4.07

###5.06###12.55###5.24###4.99###6.06###3.79

Maximal height of the roost (m)###11.31###13.95###11.18###11.71###13.70###8.39

###6.99###19.84###7.62###5.39###7.27###5.72

Maximal width of the roost (m)###10.03###12.92###9.18###12.56###11.30###8.46

###7.02###17.23###7.41###5.57###7.58###6.35

Average height of all roosts (m)###7.16###11.63###6.77###8.35###8.04###6.10

###3.66###18.38###3.38###4.63###2.939###4.33

Average weigh of all roosts (m)###6.76###9.26###5.86###9.46###6.91###6.56

###3.89###12.52###3.350###4.55###3.23###4.78

Total length of all roosts (m)###511.85###394.06###599.33###166.00###639.09###311.67

###544.26###539.09###526.12###72.66###472.30###221.58

Air temperature (C)###20.54###21.89###20.15###21.72###21.11###18.74

###3.59###3.24###3.12###4.99###1.85###1.97

Relative humidity in roost (%)###84.47###78.38###84.06###85.70###87.35###81.47

###5.12###12.23###5.40###4.48###3.08###4.26

Total floor area covered by water (m2)###26.10###31.147###29.00###17.40###23.09###30.33

###28.14###31.11###31.04###16.21###25.96###31.24

Volume of the roost (m3)###21503.42###189014.35###24613.53###12869.07###31661.32###8639.61

###28312.65###842229.52###31255.79###7193.27###35280.00###8639.08

Light intensity (lux)###6.08###6.70###0.10###24.04###0.10.###13.94

###16.95###21.74###0.00###28.76###00###23.65

the hibernacula had a lower light intensity. Moreover in a comparison of the 11 breeding roosts of H. armiger with the nine non-breeding roosts the results indicate that there was a significant difference in relative humidity and temperature between the roosts (pless than 0.05). Breeding roosts had higher air temperature and relative humidity (Table ).

For the 20 roosts used by H. armiger cave entrances facing north and northeast account for 20% each and cave entrances facing south northwest and southwest account for 13.3% each.

Finally cave entrances facing east west and southeast account for 6.7% each. Among the 34 roosts inhabited by other species cave entrances facing north account for 20%. Cave entrances facing south west and northeast account for 16% each. Cave entrances facing east and northwest account for 4% each. Cave entrances facing southeast and southwest account for 12% each (Fig. 2a). According to the Chi-square test there was no significant difference in the orientation of the entrances between the 20 roosts inhabited by H. armiger and the 34 roosts not inhabited by this there are four different types of caves: 51 (73.9%) natural caves 8 (11.6%) abandoned mines 2 (2.9%) water channels and 8 (11.6%) air-raid shelters. For the 20 roosts used by H. armiger natural caves accounted for 70% abandoned mines accounted for 20% and water channels and air-raid shelters accounted for 5% each.

Among the 34 roosts used by other species 70.6% were natural caves 8.8% were abandoned mines 2.9% were water channels and the remaining 17.7% were air-raid shelters (Fig. 2 b). According to the Chi-square test there was no significant difference in the number of the cave types between the 20 roosts inhabited by H. armiger and the 34 roosts inhabited by other species and not H. armiger (x = 2.912 df = 3 p = 0.405).

Degree of disturbance

When the roosts used by H. armiger including both the hibernation and breeding roosts were compared with non-hibernacula and non- breeding roosts there was no significant difference in the distance to the nearest road or building (Table ) but this distance was related to the degree of disturbance. The distance to the roosts under serious disturbance was smallest whereas roosts in the nature state had a longer distance to areas of human disturbance. Among the 54 roosts inhabited by bats 14 roosts were under serious disturbance 18 roosts were under light disturbance and 22 roosts were in natural state. When the 20 roosts inhabited by H. armiger were compared with 34 roosts not occupied by this species a significant difference was found in the degree of disturbance (p less than 0.05). Roosts inhabited by H. armiger had relatively less disturbance (Table ). Many H. armiger in 7 roosts were hunted by people leading to a sharp reduction in the number of them.

Other species were also hunted by people as food. Principal component analyses were performed on the variables of the 20 roosts inhabited by H. armiger. The first four principal components account for 80.58% of the total variation which best reflects the roost selection of H. armiger. The largest absolute load values of the first principal components are observed for the maximal height of the roost and the maximal width of the roost. The largest absolute load values of the second principal components are the total lengths of the roost and the volume of the roost. The largest absolute load value of the third principal components is the relative humidity in the roost. The largest absolute load value of the fourth principal components is the distance to the nearest permanent water source (Table V).

Habits

Different species of bats generally occupy

Table II.- The comparison of the characteristics and distance to the human disturbances of roosts used by H. armiger.

###Relative###Disturbance

###Number of###Temperature###Total lengths###Volume of###Light

###humidity in###distance of

###entrances###in roosts###of all tunnels###roosts###intensity

###roosts###roosts

HR and OR

T value###0.480###2.55###-1.415###0.773###-0.886###-0.109###0.072

P###0.633###0.014###0.163###0.443###0.380###0.914###0.943

HRW and HRS

T value###-2.165###-0.610###-0.836###1.804###0.819###-3.418###2.087

P###0.09###0.550###0.414###0.088###0.424###0.03###0.055

HB and HNB

T value###0.694###3.574###2.774###1.908###2.116###-1.756###1.761

P###0.497###0.002###0.013###0.072###0.048###0.117###0.096

Table III.- The comparison of the degree of human disturbance between roosts used by H. armiger and roosts not used by this species.

Degree###HR###OR

of

###Count###Percentage###Count###Percentage

disturbance

SI###3###15.0###11###32.4

LI###4###20.0###14###41.2

NS###13###65.0###9###26.5

natural caves and abandoned mines H. armiger always perched on the highest point of their roosts 30 to 150 m from the entrances in summer. In winter they usually inhabited deeper and lower parts of the roost. For human made air-raid shelters and water channels the roost sites of H. armiger were close to the entrances in summer. Nevertheless in winter H. armiger usually inhabited the middle of channels 200 m away from the entrances (Fig. 3). They usually form a colony and perch together on the inner recesses of the smooth ceiling where there is less light. H. armiger do not typically co-inhabit roosts with other species. However sometimes they co-inhabit roosts with Hipposideros pratti.

The different positions within the roosting caves. For colony size of H. armiger usually ranges from

Table V.- Rotated component matrix on the loading coefficients of morphometric data for the roosts.

###Components

###1###2###3###4

Number of entrances###0.276###0.678###-0.002###-0.362

Distance to nearest

###0.366###0.152###-0.440###0.638

permanent water (m)

Maximal height of

###0.847###0.054###0.013###-0.408

the roost (m)

Maximal width of

###0.903###0.030###0.169###-0.057

the roost (m)

Total length of all

###-0.416###0.740###0.434###0.190

roosts (m)

Air temperature (C)###0.154###-0.219###0.632###0.478

Relative humidity in

###0.310###0.119###-0.722###0.381

roost (%)

Volume of the roost

###0.239###0.839###0.280###0.235

(m3)

Light intensity (lux)###0.433###-0.488###0.550###0.218

several to hundreds of individuals and bats assemble with individual distances of approximately 100-150 mm from each other.

Pregnant females give birth to a single young each year between May and early June. A multitude of male and female H. armiger roost in separate caves during parturition and the nursing period and only a minority of males stay within maternity colonies during the breeding season.

H. armiger is less sensitive to light stimulation than other small bat species. In the caves with a light intensity of 0 less than 0.1 lux when bat species were stimulated by a torch light small bat species flew away within 4 s but H. armiger scattered after 10 s. If the height of the site used by H. armiger was more than 25 m high only a few bats flew away the rest did not leave.

H. armiger inhabited caves during the day in the summer and autumn months. Sometimes they flew around in the caves and did not fly out of the caves. They started cross-flying in caves at approximately 19:30 and a few started flying out of the cave 5 to 10 min later. After half an hour a flock of H. armiger together with other species flew out. All individuals had flown out by approximately 22:00. The first bat returned to roost at approximately 4:00. Then the bats flew back constantly until daybreak. All individuals returned to the roost site at approximately 6:00. After cross- flying for 5 to 10 min they started perching on suitable sites in different flock sizes. H. armiger leave the roost to forage in light rain but would not fly out in heavy rain.

H. armiger begin to hibernate in caves from early November until April of the following year. The period of hibernation is approximately five months. When the temperature fell to 13C in early November a portion of the H. armiger moved from the original roost which had several entrances short tunnels and unstable temperatures to a more suitable roost. H. armiger occupying roosts with single holes long tunnels and stable temperatures will moved deeper into the cave to hibernate.

DISCUSSION

In mainland China H. armiger lives in tropical and subtropical areas. These areas have higher temperatures and humidity which may be the main conditions required for the survival of H. armiger. They inhabit areas with elevation less than 2100 m above sea level.

Vonhof and Barclay (1996) suggested that some forest-dwelling bats prefer to roost on taller trees to avoid potential terrestrial predators e.g. weasels. Thus roosting on a higher ceiling may also help reduce the energy cost of staying alert or responding to the occasional disturbances and it may be beneficial for avoiding terrestrial predators. Larger spaces may provide more space and various microclimates in which bats can roost but they may also create a problem with dissipating heat and providing less total insulation for bats (Kurta 1985). In our study the roost caves used by bats have a larger volume and length than those that are unused. There were no significant differences in the total lengths and the maximal heights of the roosts between the roosts occupied by H. armiger and those used by other bat species. These results are inconsistent with those of Ho and Lee (2003).

However similar to the findings of Ho and Lee (2003) H. armiger always roost at the highest and widest positions of the cave and use deeper caves for breeding and hibernating.

The average body weight and body length of an adult H. armiger are approximately 55-70 g and 90-110 mm respectively. Because of their larger body size and presumably lower basal metabolic rate they generally maintain individual distances of 150 mm when roosting instead of clustering together. The problem of heat loss in large caves may not be as critical for this species.

Furthermore during the breeding and young rearing periods the males and females of H. armiger usually live in different roosts except for a few males that remain with the females in the breeding place which indicates that potential variations may exist in the population structure and life cycles among different colonies.

Bats usually have a higher rate of heat and evaporative water loss due to their relatively high surface area-to-volume ratio. Plecofus auritus can lose 20% to 30% of its body mass via evaporative water loss (Webb et al. 1995). Replenishing water after daily torpor is thus important for bats. Eptesicus fuscus and P. auritus select roosts that are closer to water (Entwistle et al. 1997; Williams and Brittingham 1997). In our study the roosts used by bats had higher temperatures and humidity than the unused roosts. However there was no significant difference in the distance to the nearest permanent water source between the roosts occupied by H. armiger and those used by other bat species. For the 20 roosts occupied by H. armiger the distance to the nearest permanent water was within 500 m. Because of the strong flying ability of H. armiger this distance is insignificant. A similar scenario was also proposed by Jenkins et al. (1998).

Another factor relevant to evaporative water loss is the humidity in the roosts. Bats tend to select roosts with high relative humidity (Herreid 1963; Clawson et al. 1980; Churchill 1991; Clark et al. 1996). Webb et al. (1995) found that high ambient temperatures and relative humidity would reduce the rate of the evaporative water loss of active bats. However in our study obvious differences in the humidity and temperature only exist between the breeding and non-breeding roosts of H. armiger and the breeding roosts had a higher relative humidity and temperature. Thus the humidity and temperature requirements of lactating H. armiger are higher.

Human disturbance is a major threat to the survival of many bat species and it may influence the bats' roosting behavior and roost site selection (Speakman et al. 1991). In conclusion H. armiger is selective regarding its roosts. It prefers roosts with higher ceilings larger spaces high relative humidity and little human disturbance.

H. armiger and other bat species are threatened and their populations are decreasing sharply (Zhang et al. 2009). We recommend the following measures for protecting these species: (1) develop good education programs to educate all levels of society and to raise their awareness about wildlife protection. (2) strengthen the legal protection of bats and their roosts and (3) because the best way to save a species is to protect their habitats the establishment of nature reserves should be strengthened.

ACKNOWLEDGEMENTS

This project was supported by the National Natural Science Foundation of China (NSFC No.31172056 31172050 31372163). We thank all those who helped in the field especially Songqiang Zhao Wenzhi Yang Yankun Zhu Wei Liu Yanxiao Wang and Xiao Sun. We thank Qiyun Yin Mingguo Li Junhui Chen Yumei Xiao Yudao Xu Youqiang Zheng Yulai Huang Feng Xiang for help in measure roost parameters and Xinping He Dr. Jinyou Ma for help in identification of species and Dr. Xiaojin Zhao Dr. Lina Jiang Dr. Yun Shao for help in statistical advice.

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Publication:Pakistan Journal of Zoology
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