zhu et al, 2012 (morphology, echolocation calls and diet of scotophikus kuhlii (chiroptera,...
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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: [email protected]
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.
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Received 25 October 2011, accepted 30 April 2012