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Morphology, Echolocation Calls and Diet of Scotophilus kuhlii (Chiroptera:Vespertilionidae) on Hainan Island, South ChinaAuthor(s): Guangjian Zhu, Aleksei Chmura, and Libiao ZhangSource: Acta Chiropterologica, 14(1):175-181. 2012.Published By: Museum and Institute of Zoology, Polish Academy of SciencesDOI: http://dx.doi.org/10.3161/150811012X654394URL: http://www.bioone.org/doi/full/10.3161/150811012X654394

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INTRODUCTION

Wing morphology and echolocation calls, and

their relationship to foraging ecology, are often

studied as indicators of potential resource partition-

ing by insectivorous bats (Aldridge and Rautenbach,

1987; Jones et al., 1993; Fenton and Bogdanowicz,

2002; Jennings et al., 2004; Salsamendi et al.,2005). Wing morphology strongly affects the flight

ability of bats (Norberg and Rayner, 1987) and is

often quantified using the metrics of wing load-

ing, wing aspect ratio and wing-tip shape index.

Wing loading is considered to be positively corre-

lated with minimum speed and negatively with

maneuverability. High wing aspect ratios corre-

spond to long narrow wings and energy-efficient

flight, while wing-tip shape index describes the

‘pointedness’ of wing tips and correlates with the

hovering ability in bats (Aldridge and Rautenbach,

1987; Norberg and Rayner, 1987; Jennings et al.,2004).

Insectivorous bats use echolocation calls for ob-

ject localization (e.g., prey) and for orientation in

three-dimensional space (Griffin et al., 1960). The

structure of echolocation calls reflects the degree of

acoustic clutter encountered by the bat and is indica-

tive of the size and type of prey taken (Schnitzler etal., 2003). Bats emitting loud, low frequency

echolocation calls are able to detect prey over great

distances but for the large prey only (Jennings et al.,2004). Bats alter the structure of their calls during

foraging in different habitats to efficiently hunt prey

(Schnitzler et al., 2003). In cluttered environments,

the interval between pulses is reduced since bats

need to obtain more information about the position

of obstacles to rapidly adjust their flight (Findley

and Black, 1983). Similarly, calls also tend to be re-

duced in duration as bats enter clutter to minimize

pulse-echo overlap (Kalko and Schnitzler, 1993).

The house bat, genus Scotophilus, consists of

13 species worldwide, ranging from Southeast Asia

to Africa. In China, only two species (S. kuhlii and

Acta Chiropterologica, 14(1): 175–181, 2012PL ISSN 1508-1109 © Museum and Institute of Zoology PAS

doi: 10.3161/150811012X654394

Morphology, echolocation calls and diet of Scotophilus kuhlii(Chiroptera: Vespertilionidae) on Hainan Island, south China

GUANGJIAN ZHU1, 3, ALEKSEI CHMURA2, and LIBIAO ZHANG3, 4

1Institute of Molecular Ecology and Evolution, Institute for Advanced Interdisciplinary Research, East China Normal University,Zhongshan Bei Road, Putuo Distriction, Shanghai, 200062, China

2EcoHealth Alliance, 460 West 34th Street, 17th Floor, New York, NY 10001, USA3Guangdong Entomological Institute, 105 Xingang Xi Road, Haizhu Distriction, Guangzhou, 510260, China

4Corresponding author: E-mail: zhanglb@gdei.gd.cn

Scotophilus kuhlii is distributed in many urban environments, yet the ecology of this species is poorly known. The morphology,

echolocation call structure, diet, and foraging areas of S. kuhlii were studied on Hainan Island, south China from March to

November 2006. Data from 85 individuals indicate that S. kuhlii is a medium-sized bat with 50.41 ± 1.36 (0 ± SE) mm forearm

length and 19.81 ± 3.47 g body mass. The wing morphology with high wing-loading (11.38 ± 1.95 N/m2) and moderate aspect

ratio (6.96 ± 0.75) indicates that S. kuhlii flies fast and forages in open habitat and at the edges of cluttered environments.

Echolocation calls of S. kuhlii consist of a fundamental and up to four harmonics, with a dominant frequency of 45.72 ± 2.09 kHz,

and call shape suggests that this species is adapted to forage in open environments. Data from mist-netting and acoustic detection

indicated that S. kuhlii foraged mainly around the crown of trees and street lights. Nine insect orders were recorded in its diet,

with Lepidoptera (97.46%, by frequency) and Coleoptera (64.72 ± 2.37%, by volume) constituting the main prey, together with

Hemiptera (19.99 ± 1.25%) and Hymenoptera (9.43 ± 1.14%). There was significant seasonal variation in the diet of S. kuhlii:Coleoptera increased from March to May, and then decreased to August, while Hemiptera and Hymenoptera showed the inverse

trend.

Key words: Hainan Island, bats, echolocation, morphology, ecology, diet, Scotophilus kuhlii

S. heathi) have been recorded. Scotophilus kuhlii is

found primarily in south-east Asia (Simmons, 2005)

and very little is known about its behaviour or ecol-

ogy. This species uses leaf tents as day roosting site

in the Philippines (Rickart et al., 1989; Bates and

Har rison, 1997). In this paper, we describe the mor-

phology, echolocation calls, foraging behaviour,

and seasonal variation of the diet of S. kuhlii from

Hainan Island, to better understand the natural histo-

ry of this species.

MATERIALS AND METHODS

Study Area and Sampling

Eight colonies of S. kuhlii on the campus of Hainan Normal

University, Haikou, Hainan Island, south China (19°59.910’N,

110°20.399’E) were studied. The roosts were located in a busy

urban area, with the vegetation around the roost sites consisting

of Cocos nucifera, Ficus microcarpa, Terminalia catappa and

shrubbery.

Morphological Measurements

Individuals of S. kuhlii were caught using a hand net at-

tached to a telescopic pole. For each individual, we recorded the

sex, measured body mass using a digital pocket scale (Shen -

Zhen Viabetter Electronic Scale Limited, HF-07; accurate to

0.01 g), and took the following measurements using vernier cal-

lipers (Shanghai Everwin Tool Co., Ltd; accuracy 0.01 mm):

length of the right forearm, right wing, ear, tragus, body (from

the tip of the snout to the anus ventrally) and hind foot. The out-

line of the right wing (including the right tail membrane and

the right half of the body) was traced onto paper and scanned

into a computer with same format, calculated in Photoshop 7.0

software. Wing loading, aspect ratio and wing-tip shape index

were calculated later according to Norberg and Rayner (1987).

Echolocation Call Recording

Echolocation calls were recorded from individual bats re-

leased at dusk from the centre of a soccer field located near the

roost sites. Time expansion (10X) recordings were made with

a Pettersson D-980 bat detector and digitized onto a laptop com-

puter with a sampling rate of 44.1 kHz with 16-bit precision.

BatSound Pro v3.31 (Pettersson Elektronik AB, Sweden), was

used to generate spectrograms and power spectra using a 1024-

point FFT (with Hanning window), giving 342 Hz resolution.

The dominant frequency was measured from the power spec-

trum. Start and end frequencies of the fundamental were

obtained from the oscillogram in combined Hanning windows.

Call duration and interpulse interval were measured from the

oscillogram. Interpulse interval was measured from the begin-

ning of one call to the beginning of the subsequent call.

Foraging Area Investigation

Foraging areas were determined by mist-netting, acoustic

detection (with bat detectors) and visual observations. Foraging

habitats were classified as 1) around street lights, mainly in

a street about two km from roosts, 2) around the crowns of the

trees (Ficus spp. and C. nucifera) covering an area of nearly

2 km2, 3) open areas — two sites with an estimated total area of

3 km2 , a grassland surrounding with C. nucifera and 4) over

water — Hongchen Lake approximately 0.25 km2 located in the

centre of Haikou city. Based on previous observations, four

groups (one charged with recording at fixed sites while another

checked mist-net in each group) were set in each habitat to

check mist-nets and monitored bat feeding buzzes with a Pet -

ters son D-980 bat detector during netting nights. Recordings

were analysed in BatSound Pro v3.31 (Pettersson Elektronik

AB, Sweden) software packages. Three U-shape mist-nets

(20 meters long and 10 meters high, consisted of nine single

mist-nets) were set every 500 meters at the 12 meter height in

each habitat. Every mist-net were checked every two hours dur-

ing a netting night (6:00 PM till 6:00 AM). Each habitat was

sampled for 14 consecutive nights to determine the foraging ar-

eas. The identity of flying bats was confirmed for netted indi-

viduals, and all the netted bats were marked with a 3.5 mm alu-

minium alloy bat ring (Porzana, UK) on right forearm to avoid

pseudo replication. While mist-netting, feeding buzzes were

monitored using bat detectors to determine feeding activity.

Diet Analysis

Faecal pellets were collected between March and November

2006 from roosted under folded-down leaves, commonly known

as tents, of C. nucifera (coco palms). No samples were collect-

ed between December and February as bats were absent from

the site during this period (a minimum of 60 pellets each

month). Faeces were collected from a plastic sheet placed under

each of the eight roosts for three consecutive days in the middle

of each month. Intact faeces were placed into two ml tubes con-

taining 75% ethanol and analyzed using the method described

by Kunz and Whitaker (1983). Insect remains in the droppings

were identified to order under a dissecting microscope using

the key of Zheng and Gui (1999). The percentage volume of

insect orders in each faecal sample was estimated by eye.

For Lepi doptera, only appearance frequency was recorded as it

is very difficult to estimate the percentage volume (Kunz and

Whitaker, 1983).

Statistical Analysis

All data were analyzed using SPSS 13.0 (SPSS Inc, USA).

Firstly, data normality and homoscedasticity were tested

(P > 0.05). Then, the Student’s t-test for independent samples

was used to determine if there were any differences in morphol-

ogy and echolocation call parameters between male and female.

Seasonal variation in the diet was compared using a Kruskal-

Wallis H-test, and Chi-squared test to determine the distribution

of foraging area selectivity.

RESULTS

Morphology

Eighty five adult S. kuhlii (35 XX and 50 YY)

from roosts in coco palms were captured. Morphol -

ogically, females had significantly longer forearm

(XX: 51.3 ± 1.57 mm, YY: 49.66 ± 1.07 mm;

176 G. Zhu, A. Chmura, and L. Zhang

t = 5.30, d.f. = 83, P < 0.001) and larger hand-wing

area (t = 1.15, d.f. = 83, P < 0.001) than those of

males, but no other sexual dimorphism in body

size or wing morphology was observed. S. kuhlii isa medium-sized bat with forearm length (50.41 ±

1.36 mm) and body mass (19.81 ± 3.47 g), with

a high wing-loading (11.38 ± 1.95 N/m2), moderate

aspect ratio (6.96 ± 0.75) and high tip shape index

(1.30 ± 0.52) (Table 1).

Echolocation Calls

Echolocation calls for S. kuhlii had a fundamen-

tal and up to four harmonics (Fig. 1). Each call be-

gins with a steep frequency-modulated section

and ends with a Quasi Constant Frequency part. The

frequency with most energy (dominant frequency)

was always in the fundamental. Independent-sample

t-tests showed no significant difference between

calls from males and females or any other parame-

ters measured; therefore data of echolocation calls

were combined from both sexes for subsequent

analyses. The fundamental harmonic has a starting

frequency of 89.07 ± 7.25 kHz, a dominant frequen-

cy of 45.72 ± 2.09 kHz, and an end frequency of

37.95 ± 4.05 kHz. The pulse duration and pulse

interval were 5.27 ± 1.21 ms and 44.72 ± 7.50 ms,

respectively (Table 1).

Foraging Area and Diet

One hundred and two individuals were netted

during the 14 netting nights. Most (47.1%) were

caught near the crowns of trees, 31.4% around street

lights, 16.7% above the soccer field, and 4.9% over

the lake. Frequencies of captured bats indicated

a significant difference in the use of foraging hab -

itats (χ2 = 40.82, d.f. = 3, P < 0.001), indicating that

S. kuhlii mainly foraged near the crowns of trees and

around the street lights. While, 12, 3, 1 and 0 feed-

ing buzzes were recorded during 14 nights bat

detecting in the four habitat types, crown of trees,

around street lights, soccer field and above the lake

respectively. This supports the results from mist-

netting data (Table 2).

Nine insect orders were identified from the fae-

ces of S. kuhlii. By volume, in decreasing order

were Coleoptera (64.72 ± 2.37%), Hemiptera

(19.99 ± 1.25%), Hymenoptera (9.43 ± 1.14%), and

Diptera (5.49 ± 0.14%), while Lepidoptera (97.46%)

were recorded the highest by frequency. Odonata,

Homo ptera, Trichoptera, Orthoptera were occasion-

ally recorded in the faeces (Table 3).

The main prey items taken by S. kuhlii showed

significant seasonal variation (Lepidoptera: χ2 = 39.4,

d.f. = 9, P < 0.001; Coleoptera: χ2 = 402.7, d.f. = 9,

P < 0.001; Hemiptera: χ2 = 268.0, d.f. = 9, P < 0.001;

Morphology, echolocation calls and diet of Scotophilus kuhlii 177

Parameter0 ± SE Range

PXX YY XX YY

Body size Forearm length (mm) 51.3 ± 1.6 49.7 ± 1.1 48.1–54.6 48.0–51.9 0.000

Body mass (g) 20.5 ± 3.6 19.1 ± 3.8 12.7–29.3 12.5–30.3 0.091

Ear length (mm) 10.5 ± 1.1 10.5 ± 1.2 7.5–12.4 8.3–13.4 0.846

Tragus length (mm) 6.3 ± 0.6 6.3 ± 0.7 5.1–7.4 5.1–7.2 0.707

Hind foot length (mm) 10.4 ± 0.8 10.2 ± 1.2 8.1–12.1 8.3–13.8 0.448

Body length (mm) 69.5 ± 5.6 67.2 ± 5.4 54.8–81.3 54.8–82.1 0.056

Tail length (mm) 42.3 ± 4.4 40.2 ± 4.3 31.3–49.3 31.5–50.1 0.069

Wing morphology Wing span (mm) 34.9 ± 1.9 33.9 ± 1.5 28.8–38.2 30.4–36.6 0.060

Wing area (×10-3 m2) 17.7 ± 2.1 16.6 ± 1.9 10.7–21.9 12.8–20.1 0.021

Aspect ratio 7.0 ± 0.7 7.0 ± 0.8 5.6–8.7 5.3–10.0 0.954

Wing loading (N/m2) 11.5 ± 2.1 11.3 ± 1.9 7.0–17.5 8.0–16.7 0.704

Hand-wing area (×10-3 m2) 3.1 ± 0.3 2.8 ± 0.3 2.3–3.8 2.2–3.4 0.000

Arm-wing area (×10-3 m2) 4.1 ± 0.6 4.0 ± 0.6 3.1–5.6 2.7–5.0 0.232

Tip length ratio 1.4 ± 0.1 1.4 ± 0.1 1.1–1.7 1.1–1.5 0.454

Tip area ratio 0.8 ± 0.9 0.8 ± 1.0 0.6–1.0 0.5–1.0 0.110

Tip shape index 1.4 ± 0.7 1.3 ± 0.5 0.8–3.6 0.6–2.8 0.441

Echolocation calls Dominant frequency (kHz) 45.7 ± 2.1 45.8 ± 2.2 41.9–51.4 39.9–50.2 0.870

Start frequency (kHz) 88.7 ± 6.9 89.4 ± 7.9 64.2–103.2 71.0–104.5 0.780

End frequency (kHz) 37.9 ± 2.8 38.1 ± 5.2 31.0–44.1 12.8–42.7 0.840

Pulse duration (ms) 5.1 ± 1.0 5.4 ± 2.1 2.9–8.1 3.0–14.1 0.490

Pulse interval (ms) 46.8 ± 24.7 43.0 ± 25.2 13.5–127.4 13.1–127.4 0.530

TABLE 1. The measurements of body size, wing morphology and echolocation calls from S. kuhlii on Hainan Island,

south China. Data were obtained from 85 individuals (35 YY and 50 XX) captured in October 2006. A Student’s t-test was

conducted on all the measured characters between female and male S. kuhlii

178 G. Zhu, A. Chmura, and L. Zhang

FIG. 1. Two sequential echolocation calls recorded from S. kuhlii in free flight after release from the hand. The three panels show

the waveform (A), spectrogram (B) and power spectra (left call = C, right call = D)

Am

plit

ude (

%)

Fre

quency (

kH

z)

Pow

er

(dB

)P

ow

er

(dB

)

Frequency (kHz)

A

B

C

D

Morphology, echolocation calls and diet of Scotophilus kuhlii 179

Hymenoptera: χ2 = 244.9, d.f. = 9, P < 0.001) (Table

3). The percent volume for Coleoptera increased

from March to May, and then decreased towards

August, while Hemi ptera and Hymenoptera showed

an inverse trend.

DISCUSSION

Morphology

This study showed that there were no significant

morphological differences between male and female

S. kuhlii except forearm length and hand-wing area.

The morphological data is consistent with prior

studies conducted outside of China (Goodman et al.,2005), except ear and hind foot lengths recorded

were shorter. The average wing span of male S. kuh-lii in this study is consistent with that recorded by

Goodwin (1979). However, owing to the absence of

measurements for females (Goodwin, 1979), no

comparisons can be made for the wing span of fe-

male S. kuhlii. Due to lack of recorded data for body

mass and wing area from Goodwin (1979) and lim-

ited sample size, little can be said about differences

in wing loading and aspect ratio. Studies of other

insectivorous bats have suggested that species

with low wing loading and aspect ratio have greater

manoeuvrability (Jennings et al., 2004). This

study indicates that S. kuhlii on Hainan Island has

a high wing-loading (11.6 N/m2) and a moderate

aspect ratio (6.96). This implies an ability to fly

fast with low energy expenditure, but low manoeu-

vrability, indicating that S. kuhlii may fly and forage

in open habitat or at the edge of cluttered environ-

ment. The present study found that S. kuhlii pre-

ferred foraging in relatively open area or around the

street lights, supporting the predictions of the wing

morphology.

Echolocation Call

Studies on echolocation have established that

variations in the echolocation calls of bats exist at

three different levels: species, individuals within

a species, and calls of a given individual (Karry etal., 2001). Intraspecific echolocation variation be-

tween the sexes has been described for many species

(Neuweiler et al., 1987; Suga et al., 1987; Jones etal., 1992, 1993). The present study found no differ-

ence between the sexes for S. kuhlii. Several studies

have established a relationship between echoloca-

tion call structure, morphology, and foraging behav-

iour (Norberg and Rayner, 1987; Jones et al., 1993;

Jennings et al., 2004; Salsamendi et al., 2005).

Echolo cation call structure has a relationship with

body size (Jones et al., 1993; Zhang et al., 2007),

with larger species usually emitting calls with

a lower frequency and higher intensity compared to

smaller species. The production of lower frequency

and higher intensity pulses increases potential detec-

tion over long distances. The frequency with most

TABLE 2. Data from mist-netting and feeding buzzes in the

examined foraging area. Mist-netting was performed with U-

shape mist net groups, each mist-net was checked about every

two hours

HabitatsMist-netting Feeding buzzes

(individuals caught) (quantity recorded)

Around street light 32 3

Crown of the trees 48 12

Open areas 17 1

Above lake 5 0

MonthsFrequency percentage Volume percentage (0 ± SE)

Lepidoptera Coleoptera Hemiptera Hymenoptera Diptera

March 95.8 67.5 ± 0.8 22.1 ± 0.6 7.5 ± 0.5 2.0 ± 0.4

April 100.0 74.0 ± 1.0 17.3 ± 0.8 6.7 ± 0.5 2.0 ± 0.4

May 100.0 76.2 ± 0.5 13.1 ± 0.4 6.4 ± 0.3 4.3 ± 0.3

June 100.0 66.6 ± 0.6 18.1 ± 0.5 8.5 ± 0.3 6.4 ± 0.2

July 89.1 61.6 ± 0.5 23.4 ± 0.5 9.1 ± 0.3 5.8 ± 0.2

August 92.4 51.2 ± 0.8 23.2 ± 0.6 18.6 ± 0.9 7.0 ± 0.3

September 97.3 65.0 ± 1.0 18.4 ± 0.8 7.1 ± 0.3 7.5 ± 0.7

October 100.0 62.2 ± 1.0 24.7 ± 0.7 7.3 ± 0.4 5.0 ± 0.4

November 100.0 60.4 ± 1.3 26.6 ± 0.9 8.6 ± 0.7 3.9 ± 0.4

December 100.0 65.5 ± 1.8 19.6 ± 0.1 11.4 ± 0.7 3.2 ± 0.8

Total 97.5 64.7 ± 0.4 20.0 ± 0.3 9.4 ± 0.2 5.5 ± 0.1

χ2 39.4 402.7 268.0 244.9 195.0

TABLE 3. Analysis of monthly dietary components from S. kuhlii. 789 faecal pellets (over 60 samples each month) were

collected from March to November 2006. All the orders varied seasonally, all P < 0.001

180 G. Zhu, A. Chmura, and L. Zhang

energy was in the fundamental, with a low frequen-

cy of 45.72 kHz, indicating that S. kuhlii can detect

prey over long distances in open habitats, and may

catch relatively large prey. It also emitted relatively

broadband frequency-modulated echolocation calls

with the fourth harmonic up to 200 kHz during

flight, which may be well suited for obtaining

detailed information about the target (Neuweiler,

1984).

Diet and Foraging Area

Although radio telemetry was regarded as an im-

portant and effective tool for investigating the activ-

ities and foraging areas of mammals and birds, ultra-

sound detectors (McAney and Fairley, 1988), light-

tagging animals (Schofield and Morris, 1999) and

the mark-recapture method (Handley et al., 1991)

have been used to determine foraging areas of

bats. In view of the fact that very weak and highly

directional electronic signals of radio telemetry

makes them difficult to detect in the field, data from

ultrasound detectors obtained in such circumstances

might be difficult to interpret. So, we used mist-net-

ting, acoustic detection (via bat detectors) and visu-

al observations together to determine the foraging

areas of S. kuhlii bat in this study.

Aldridge and Rautenbach (1987) showed that

wing morphology and echolocation call structure

could influence foraging site selection and foraging

behaviour in insectivorous bats. Data from the pres-

ent study revealed that S. kuhlii foraged predomi-

nately in open environments or at the edge of the

cluttered environments, including the crowns of

trees within the urban environment, around street

lights, a soccer field, and over a lake, which support-

ed our predtion based on the wing morphology and

echolocation calls. The morphology of S. leuco-gaster is similar to S. kuhlii (Goodman et al., 2005)

and it preys primarily on Hemiptera and Coleoptera

(Barclay, 1985). Our data also indicated that S. kuh-lii feed mainly on Coleoptera, Hemiptera, Lepido -

ptera and Hymenoptera. The volume of Coleoptera

and Hemiptera in the diet approaches 80% every

month. The frequency of occurrence of Lepi-

doptera was used instead of volume in diet analysis

to prevent biased results (usually an overestimate

because of array of scale in Lepidoptera). It is

known that Lepidoptera insects have ability to detect

ultrasound, but they were a frequently recorded item

in S. kuhlii’s diet in our study. Although the best

hearing range of moth is considered to be 20–60

kHz, this varies among species. Fenton and Fullard

(1979) found that the ears of moths in Canada

are most sensitive to sounds between 20 and 40

kHz. Scotophilus kuhlii emits calls with dominate

frequency at 45 kHz, suggesting that it has the

potential ability to feed on some moth species. Al -

dridge and Rautenbach (1987) showed that echolo-

cation calls should be related to wing morphology,

and reflect the character of the prey and the foraging

habitat. The relatively low dominant frequency of

S. kuhlii echolocation calls and high wind loading

suggest a species foraging in open habitat and

feeding on relatively large prey. Lepidoptera and

Coleoptera, preyed on by S. kuhlii, are relatively

large insects.

ACKNOWLEDGEMENTS

We thank Prof. S. Y. Zhang, Prof. S. Parsons, J. S. Zhang, X.

D. Zhao, and B. Liang for their comments on earlier drafts of

this manuscript. We also thank F. Li, M. Li, J. Guilbert, and

Z. H. Tang for their help in the field. We are very grateful for

G. P. Wang’s help with the faecal analysis. This project was

supported by The Ministry of Science and Technology of the

People’s Republic of China (MOST grant no. 2006FY110500),

National Natural Science Foundation of China (NSFC, No.

30800102), and Special Foundation for Young Scientists

([2008]02) of Guangdong Province Academy of Science.

LITERATURE CITED

ALDRIDGE, H. D. J. N., and I. L. RAUTENBACH. 1987. Mor phol -

ogy, echolocation and resource partitioning in insectivorous

bats. Journal of Animal Ecology, 56: 763–778.

BARCLAY, R. M. R. 1985. Foraging behavior of the African

insectivorous bat, Scotophilus leucogaster. Biotropica, 17:

65–70.

BATES, P. J. J., and D. L. HARRISON. 1997. Bats of the Indian

subcontinent. Harrison Zoological Museum, Sevenoaks,

Kent, 258 pp.

FENTON, M. B., and W. BOGDANOWICZ. 2002. Relationships

between external morphology and foraging behaviour:

bats in the genus Myotis. Canadian Journal of Zoology, 80:

1004–1013.

FENTON, M. B., and J. H. FULLARD. 1979. The influence of moth

hearing on bat echolocation strategries. Journal of Com -

para tive physiology, 132A: 77–86.

FINDLEY, J. S., and H. BLACK. 1983. Morphological and die-

tary structuring of a Zambian insectivorous bat community.

Ecol ogy, 64: 625–630.

GOODMAN, S. M., R. K. B. JENKINS, and F. H. RATRIMOMANA -

RIVO. 2005. A review of the genus Scotophilus (Mammalia,

Chiroptera, Vespertilionidae) on Madagascar, with the de-

scription of a new species. Zoosystema, 27: 867–882.

GOODWIN, R. E. 1979. The bats of Timor: systematics and ecol-

ogy. Bulletin of the American Museum of Natural History,

163: 73–122.

GRIFFIN, D. R., F. A. WEBSTER, and C. R. MICHAEL. 1960. The

echolocation of flying insects by bats. Animal behaviour, 8:

141–154.

HANDLEY, C. O., JR., A. L. GARDNER, and D. E. WILSON. 1991.

Move ments. Pp. 89–130, in Demography and nature histo-

ry of the common fruit bat Artibeus jamaicensis on Bar-

ro Colorado Island, Panama (C. O. HANDLEY, JR., D. E.

WILSON, and A. L. GARDNER, eds.). Smithsonian Institution

Press, Wash ington D.C., 173 pp.

JENNINGS, N. V., S. PARSONS, K. E. BARLOW, and M. R. GANNON.

2004. Echolocation calls and wing morphology of bats from

the West Indies. Acta Chiropterologica, 6: 75–90.

JONES, G., T. GORDON, and J. NIGHTINGALE. 1992. Sex and age

differences in the echolocation calls of the lesser horseshoe

bat, Rhinolophus hipposideros. Mammalia, 56: 189–193.

JONES, G., M. MORTON, P. M. HUGHES, and R. M. BUDDEN.

1993. Echolocation, flight morphology and foraging strat-

egies of some West African hipposiderid bats. Journal of

Zool ogy (London), 230: 385–400.

KALKO, E. K. V., and H.-U. SCHNITZLER. 1993. Plasticity in

echolocation signals of European pipistrelle bats in search

flight — implications for habitat use and prey detection.

Behav ioral Ecology and Sociobiology, 33: 415–428.

KARRY, A. K., C. B. STEPHEN, and W. M. MASTERS. 2001.

Individual and group variation in echolocation calls of big

brown bats, Eptesicus fuscus (Chiroptera: Vespertilionidae).

Journal of Mammalogy, 82: 339–351.

KUNZ, T. H., and J. O. WHITAKER, JR. 1983. An evaluation of fe-

cal analysis for determining food habits of insectivorous

bats. Canadian Journal of Zoology, 61: 1317–1321.

MCANEY, C. M., and J. S. FAIRLEY. 1988. Habitat preference

and overnight and seasonal variation in the foraging activi-

ty of lesser horseshoe bats. Acta Theriologica, 33: 393–402.

NEUWEILER, G. 1984. Foraging, echolocation and audition in

bats. Naturwissenschaften, 71: 446–455.

NEUWEILER, G., W. METZNER, U. HEILMANN, R. RUBSAMEN,

M. ECKRICH, and H. H. COSTA. 1987. Foraging behavior and

echolocation in the rufous horseshoe bat (Rhinolophusrouxi) of Sri Lanka. Behavioral Ecology and Sociobiology,

20: 53–67.

NORBERG, U. M., and J. M. V. RAYNER. 1987. Ecological mor-

phology and flight in bats (Mammalia; Chiroptera): wing

adaptations, flight performance, foraging strategy and echo -

location. Philosophical Transactions of the Royal Society of

London, 316B: 335–427.

RICKART, E. A., P. D. HEIDEMAN, and R. C. B. UTZURRUM. 1989.

Tent-roosting by Scotophilus kuhlii (Chiroptera: Vesperti-

lio nidae) in the Philippines. Journal of Tropical Ecology, 5:

433–436.

SALSAMENDI, E., J. AIHARTZA, U. GOITI, D. ALMENAR, and

I. GARIN. 2005. Echolocation call and morphology in the

Me helyi’s (Rhinolophus mehelyi) and Mediterranean

(R. eury a le) horseshoe bats: implication for resource parti-

tioning. Hystrix, 16: 149–158.

SCHNITZLER, H-U., C. F. MOSS, and A. DENZINGER. 2003. From

spatial orientation to food acquisition in echolocating bats.

Trends in Ecology and Evolution, 18: 386–394.

SCHOFIELD, H., and C. MORRIS. 1999. The micro-habitat prefer-

ences of Bechsterin’s bat within woodlands in southern

England. Bat Research News, 40: 104–141.

SIMMONS, N. B. 2005. Order Chiroptera. Pp. 465–467, in Mam -

mal species of the World: a taxonomic and geographic ref-

erence (D. E. WILSON and D. M. REEDER, eds.). The Johns

Hopkins University Press, Baltimore, 2142 pp.

SUGA, N., H. NIWA, I. TANIGUCHI, and D. MARGOLIASH. 1987.

The personalized auditory cortex of the mustached bat:

adaptation for echolocation. Journal of Neurophysiology,

58: 643–654.

ZHANG, L., B. LIANG, P. STUART, L. WEI, and S. ZHANG.

2007. Morphology, echolocation and foraging behaviour in

two sympatric sibling bats, Tylonycteris pachypus and T. ro-bustula (Chiroptera: Vespertilionidae). Journal of Zoology

(London), 271: 344–351.

ZHENG, L., and H. GUI. 1999. Insect classification. Nanjing Nor -

mal University Press, Nanjing, Volumes 1 and 2, 1070 pp.

Morphology, echolocation calls and diet of Scotophilus kuhlii 181

Received 25 October 2011, accepted 30 April 2012