Aldridge y Rautenbach 1987 Morphology Ecolocacion and Food Partitioning in Insectivorous Bats

Morphology, Echolocation and Resource Partitioning in Insectivorous Bats
Author(s): H. D. J. N. Aldridge and I. L. Rautenbach

Source: Journal of Animal Ecology, Vol. 56, No. 3 (Oct., 1987), pp. 763-778
Published by: British Ecological Society
Stable URL: http://www.jstor.org/stable/4947
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Journal of Animal Ecology (1987), 56, 763-778
MORPHOLOGY, ECHOLOCATION AND RESOURCE

PARTITIONING IN INSECTIVOROUS BATS
BY H. D. J. N. ALDRIDGE* AND I. L. RAUTENBACH
Department of Biology, Carleton University, Ottawa KIS 5B6, Canada

and Department of Mammalogy, Transvaal Museum, Pretoria,
Republic of South Africa
SUMMARY
(1) Two hypotheses are tested in this paper: (i) wing morphology and echo
'design' can determine foraging site selection and foraging behaviour in bats, (ii)
echolocation-call 'design' should be compatible with wing morphology (because some
combinations of morphology and echolocation call would be maladaptive).
(2) In support of our first hypothesis, significant correlations were established between
wing morphology, echolocation call design, manoeuvrability and habitat use.
(3) In support of our second hypothesis, significant correlations were established
between those morphological parameters that improve manoeuvrability (low wingload-
ing, low aspect ratio and high wingtip shape index) and echolocation calls that are
resistant to acoustic clutter in support of our second hypothesis.
(4) There was an association between foraging habitat and diet: bats that fed in the
same habitats tended to take the same types of prey, while species foraging in different
habitats had significantly different diets.
(5) There was also a significant correlation between prey and predator size; large bats
took insects over a range of sizes while small bats fed only on small prey.
INTRODUCTION
The study of resource partitioning in bat communities has concentrated primarily on

morphology (reviewed by Findley & Wilson 1982) and more recently on echolocation
(reviewed by Neuweiler 1983), on the assumption that species' differences in these
characters reflect resource partitioning. A number of studies have indicated that there is a
significant causal relationship between wing morphology and feeding ecology. In an
African bat community, O'Shea & Vaughan (1980) found a significant positive
correlation between aspect ratio and flight height; bats with high aspect ratio wings
tended to fly high. Aldridge (1986a) found that foraging site selection in a British bat
community was related to manoeuvrability, a characteristic which in the North American
bats Myotis lucifugus and M. yumanensis was related to wingloading and wing span
(Aldridge 1986b).
A bat's ability to forage successfully within a particular habitat will also depend upon
the characteristics (frequency, band-width and duration) of its echolocation calls. The
role of echolocation-call 'design' in determining strategies of habitat use is unclear
because there are no data on the variability of call use within and between species. There
are, however, some general trends (reviewed by Neuweiler 1983): lower frequency, narrow
* Present address: Department of Biology, York University, Downsview, Toronto M3J 1P3, Canada.
763
764 Resource partitioning in bats
Fig. 1. Map showing the Pafuri area of the Kruger National Park, the positions of the netting and
trapping sites are indicated: solid circles = sites in riverine forest dominated by Ficus spp., solid
hexagons = small lakes or water-holes in relatively open bush/veldt; solid triangle = site within a
garden dominated by cultivated fruit trees (e.g. mangoes, lychees and bananas).
band-width calls are well suited for long-range detection of prey, but may result in the user
being confused by acoustic clutter; broad-band calls at higher frequency are more
resistant to clutter (Simmons & Stein 1980), but suffer more from atmospheric
attenuation (Griffin 1971). We would expect low frequency, narrow-band calls to be
associated with bats foraging in open areas and broad-band calls (or constant frequency
calls associated with Doppler-shift compensation) to be used by bats foraging in highly
cluttered habitats.
The aim of this study was to test two hypotheses.
(a) Morphology determines the selection of foraging habitats in bats, and differences in
morphology will result in the partitioning of spatial resources.
(b) Morphology and the 'design' of echolocation calls are inter-related as part of the
same adaptive complex so that particular call designs are associated with particular
strategies of habitat use, and therefore with particular morphologies.
MATERIALS AND METHODS
The animals
We caught twenty-six species of the sub-order Microchiroptera either in mist-nets or in

traps (Tuttle 1974) at a variety of sites in the Pafuri area of the Kruger National Park,
South Africa (Fig. 1) during November (the beginning of the wet season) 1985 (Table 1).
The bats were kept for 1-12 h and their morphological parameters recorded, faecal
samples collected and their abilities to fly through a series of obstacles tested; some bats
were light-tagged (Buchler 1976) and released almost immediately after capture.
Morhpology
To ensure that their alimentary tracts were empty the bats were kept for at least 1 h after
capture before being weighed. Each bat was weighed using an OHAUS C200M electronic
H. D. J. N. ALDRIDGE AND I. L. RAUTENBACH 765
TABLE 1. Morphological parameters of twenty-six sympatric insectivorous bat

species, resident in the Pafuri region
Mass Wingloading Aspect ratio Wingtip

Species n (kg) (N m-2) shape index
Rhinolophidae x S.D. x S.D. x S.D. x S.D.
Rhinolophus
R. hildebrandti (Peters) 9 0-024 0-0020 9.8 1-3 6-8 0-6 2-5 0-8
R. fumigatus (Ruppell) 6 0-013 0.0010 7.5 0-4 6-7 0-3 3-0 1-7
R. darlingi (Andersen) 1 0'011 7-8 - 63 - 1-7
R. landeri (Martin) 7 0'009 0.0010 6-7 0-3 6-1 0-3 2-6 1-2
R. swinnyi (Gough) 2 0-007 6-3 - 66 - 38 -
Hipposideridae
Hipposideros
H. caffer (Sundervall) 14 0-009 0-0007 6-3 0-5 6-4 0-4 2-8 0-6
H. commersoni (Geoffroy) 2 0-089 - 15.7 7-7 - 3-1
Nycteridae
Nycteris
N. thebaica (Geoffroy) 2 0.011 6-3 55 4.5
Vespertilionidae
Kerivoula
K. argentata (Tomes) 1 0-010 6-6 0' 0 6-1 2-0
Myotis
M. tricolor (Temminck) 2 0-016 8-2 6-2 2-2
M. bocagei (Peters) 3 0-008 0-0003 6-4 0-2 6-5 0-1 2-0 0-6
Pipistrellus
P. rueppellii (Fischer) 4 0-007 0-0007 6-8 0-6 6-3 0-6 2-0 1.0
P. rusticus (Tomes) 2 0-005 7-0 6-8 - 13
P. nanus (Peters) 5 0-004 0-0006 8-4 2-0 7.3 1-3 1-4 0-5
Eptesicus
E. hottentotus (A. Smith) 1 0-030 14-0 72 - I.
E. capensis (A. Smith) 11 0-007 0-0007 7 1 0-5 6-4 0-3 1-9 0-7
E. zuluensis (Roberts) 4 0-004 0-0005 5-7 1-0 6-4 0-4 1-8 0-3
Nycticeius
N. schlieffeni (Peters) 10 0-005 0-0005 6-7 1.0 6-9 0-4 1-6 0-4
Scotophilus
S. borbonicus (Geoffroy) 5 0-016 0-0002 10-4 1-2 7.5 0-5 1-5 0-2
S. dingani (A. Smith) 19 0-024 0-0020 12-4 09 7.3 0-4 2-0 0-6
Molossidae
Tadarida
T. midas (Sundervall) 4 0-044 0-0003 18-4 0.9 8-9 0-7 1-4 0-2
T. fulminans (Thomas) 1 0-033 20-2 - 14-3 - 1-5
T. condylura (Smith) 5 0-026 0-0003 18-0 1-6 9.1 0-9 1-7 0-4
T. pumila (Cretschmar) 1 0-013 12-6 - 8-9 - 1-2
Tadarida spp. 4 0-022 0-0002 15-5 1-0 9.4 0-2 1-4 0-2
Emballonuridae
Taphozous
T. mauritianus (Geoffroy) 2 0-026 15-4 - 10-1 - 1-1
balance (accurate to 0-1 g; Ohau

Jersey 07932, U.S.A.), and its w
recorded wing area by tracing t
paper. A number of different c
(Gaisler 1959; Vaughan 1959, 19
Chari 1976), but we followed that
measured using a Placom electronic planimeter. For each individual three areas were
recorded, the area of one wing plus one half of the areas of the body and tail, and the areas
of the handwing (the chiropatagium) and the armwing (the plagiopatagium and
propatagium; Fig. 2).
Armwing (Sp)
Tail
FIG. 2. Schematic diagram showing the morphological parameters measured in this study. Lp is
the length of the armwing, Lc the length of the handwing, Sp is the area of the armwing
(plagiopatagium and propatagium) and Sc is the area of the handwing (chiropatagium; shaded
area). Wing area is measured as the area of the wing, the tail and the body, but excluding the area
of the head.
The fundamental measurements were used to derive three further parameters:

wingloading, aspect ratio and wingtip shape index. Wingloading (Qs) is defined as:
Qs= m g/S (1)
where m is body mass, g the accel
is defined as:
R = b2/S (2)
where b is wing span.
The idea that the shape of the
lity is generally accepted (Vaughan 1959; Findley, Studier & Wilson 1972; Smith &
Starrett 1979; Findley & Wilson 1982; Aldridge 1986a, b). Several indices describing
wingtip shape have been devised (e.g. Findley, Studier & Wilson 1972; Smith & Starrett
1979), but U.M. Norberg and J.M.V. Rayner (personal communication) have shown that
these tend to be correlated with aspect ratio and therefore to describe the overall shape of
the wings. Norberg and Rayner (personal communication) have devised an index, I, that
is not highly correlated with aspect ratio but that describes the proportion that the
handwing and the armwing contribute to the total wing. I is defined as:
I= TsIT, -Ts (3)
where Ts is the ratio of the area of
ratio of the handwing length and
values, while pointed ones have low
Echolocation calls
We used a QMC S-200 bat detector with a period meter and a NLS 20 Miniscope
(Simmons et al. 1979), to monitor echolocation calls from free-flying light-tagged bats
TABLE 2. Characteristics of the search-phase echolocation calls of sixteen sympatric

insectivorous bat species resident in the Pafuri region. The lowest and highest
frequencies and the frequency at which the call is most intense are shown. sh-FM,
st-sh-FM, st-FM and CF-FM are shallow sweep frequency-modulated signals,
steep and then shallow FM signals, steep FM signals and constant frequency signals
which begin, end or both with steep FM signals, respectively. n1 is the number of
bats from which calls were observed in the flight cage and n2 the number released
with light-tags from which observations were made
Frequency
low high Duration
Species n\ n2 (kHz) (kHz) Intensity (ms) Shape
R. hildebrandti 9 7 c.24 c.40 high 15 CF-FM
R. fumigatus 6 50 55 high 15 CF-FM
R. landeri 5 - 105 110 high 15 CF-FM
R. swinnyi 1 - 100 115 med. 15 CF-FM
H. caffer 5 20 105 138 high 7 CF-FM
H. commersoni 2 - 55 62 high 12 CF-FM
N. thebaica 4 - 61 97 low 2 st-FM
P. rueppellii 5 - 40 70 high 4 st-FM
P. nanus 5 20 62 90 high 5 st-FM
E. capensis 4 20 35 65 high 5 st-FM
N. schlieffeni 8 20 33 78 high 5 st-FM
S. borbonicus 7 7 40 70 high 5 st-sh-FM
S. dingani 6 9 35 58 high 5 st-sh-FM
T. midas - ? ? 30 high 12 sh-FM
(recorded as bats left roost)
T. fulminans 1 - 14 27 high 20 sh-FM
T. mauritianus 2 - 15 59 high 20 sh-FM
(Buchler 1976). After the light-tags were attached t

they flew towards the microphone. For each speci
(kHz) and the frequency of maximum intensity were recorded directly from the bat
detector. The shapes and duration of the search phase calls were transcribed by hand from
the oscilloscope screen (Table 2). For those species for which we lacked light-tag data,
echolocation calls were monitored as the bats flew through the obstacle-free flight tunnel
(see below). Echolocation calls were characterized by their frequency and their shape: call
shape i is defined as:
A = bf/t (4)
wherefis change in call frequen
Manoeuvrability
We tested the bats' manoeuvrabilities by flying them through a series of obstacles
arranged within a flight tunnel (4-0 x 0O88 x 0-88 m), at either end of which we erected
two gauze tents (c. 2 x 2 x 2 m). The obstacles were parallel vertical assays of pieces of 3
mm diameter binder-twine arranged so that the strings were equidistant from one
another. Each bat was tested with three arrangements of the strings. The first arrangement
of the series was chosen randomly from seven possible arrangements (open tunnel and six
string arrangements with inter-string distances (d) of 0-6 m, 0-44 m, 0-3 m, 0O22 m, 0-15 m
and 0- 1 m). If the bat was unable to negotiate the first arrangement it was challenged with
the next easiest in the series. If it was able to negotiate the obstacles then it was challenged
with a more difficult arrangement. After testing a number of individuals of a given species
FIG. 3. Habitat zones used to record bat foraging behaviour: Zone 1, open areas; zone 2, within 0-5
m of water surface; zone 3, over pasture; zone 4, around trees, 0-5 m away from canopy, zone 5,
within woodland, between larger branches and trunks of trees; zone 6, the surface branches and
leaves of trees and bushes; zone 7, within canopy or thicket.
we could adjust the arrangement at which we began testing and ensure that we obtained
the most usable data from relatively small samples of each species. To minimize the
number of refusals that might result from the stress associated with the procedure, each
bat was given three tests with a particular arrangement, although in most cases (93% of
215 trials) the bats attempted to fly through the tunnel on the first test. Manoeuvrability
was scored as the minimum inter-string distance an individual could negotiate (mininum
negotiable distance, MND).
Habitat use
Potential habitat available to the bats was subdivided into seven foraging zones, ranked
according to a subjective assessment of the density of clutter within each (Fig. 3; Aldridge
1986a). The bats' use of these zones was recorded by continuously monitoring the
foraging flight of light-tagged individuals (Buchler 1976).
We used size 000 gelatine pill capsules filled with Cyalume (American Cyanamid
Company, Chemical Light Department, Milton, Florida 32570, U.S.A.), and attached
them by surgical adhesive to the bats' fur just below the junction of the wing membrane
and the abdomen. The foraging flights of the bats were described on hand-held cassette-
tape recorders. For each individual we noted its position within the foraging zones
described above and the nature of its foraging behaviour (i.e. continuous flight, sallying or
hovering and gleaning). From the tapes we recorded the duration of the bat's use of each
zone, and described its behaviour within that zone. QMC mini-bat-detectors were used to
monitor echolocation calls to distinguish between foraging and commuting flights. The
bats were always released where they had been netted.
0.
20
FIG. 4. Body mass, wingloading and aspect ratio for twenty-six sympatric microchiropteran bat
species caught in the Pafuri region. The bats were divided into four groups (for details see text).
Group 1 (0): E. zuluensis, P. nanus, P. rusticus, N. schlieffeni, P. rueppellii, C. capensis, H. caffer,
M. bocagei, R. swinnyi, N. thebaica, R. landeri, K. argentata, R. darlingi and R.fumigatus. Group 2
(a): R. hildebrandti, M. tricolor, E. hottentotus, S. borbonicus and S. dingani. Group 3 (A): H.
commersoni. Group 4 (M): Tadarida sp., T. pumila, T. condylura, T. mauritianus, T.fulminans and
T. midas.
For each individual a habitat use index (HU) was calculated using the following
equation:
7
HU = E (Ht^/) (5)
H= I
where His the rank of the zone, th the time spent in zone of rank H, and t
which the bat was observed.
Diet
Bats that had obviously fed before capture were individually housed in glass storage
jars or polystyrene cups and left for at least 1 h. We collected faeces from each bat and
analysed samples of up to ten pellets under a binocular dissecting microscope. The
droppings were teased apart under weak ethanol or detergent solutions and the insects
therein identified to order. We measured the lengths of beetle (Coleoptera) tarsal claws
and estimated the minimum number of insects that could have been ingested to produce
the observed number of insect parts. When there were no easily identifiable parts (e.g. legs,
claws, head capsules, antennae and wings) other than scales (Lepidoptera and
Trichoptera) we estimated what percentage the scales were of the total volume of the
pellet.
One of our aims was to establish whether there was any relationship between predator
and prey size. We found enough beetle claws in the pellets to measure prey size and to
compare the diets of the different species. Fortunately, we were able to measure the claw
and body lengths for thirty-six beetle species ranging in length from c. 2 mm to almost 50
160- (a) \o
80- 160-- -24
80 - \
-9 0 \,
1 40- 0 ~ 80- -
LLi
20- o\ 40- 8 \
.0 ~10
0 -I
.1 ,-~
I 0-1
I 25-
I I 2
0
5 10 20 40 3 6 12 24
Wingloading (N m-2) Aspect ratio
(c)
/
I 160- / -20
0
@ 80- a)
o 0 / -12
40- 0o/ o
25- , -0
1-5 3 6 10
Wingtip shape
FIG. 5. Double log plots showing the

frequency (only highest frequencies
circles and hatched line): (a) wingloa
mm. There was a significant relationship between claw and body length (r =0-75,
P<0.01), permitting us to estimate beetle size from claw size. We did not use the
numerical estimates of prey size in the diet analysis, but assigned the beetles to three size
classes: large, beetles 20 mm; medium, 10-20 mm, and small, 10 mm.
RESULTS
Manoeuvrability and habitat-use predictions

We obtained morphological data for twenty-six microchiropteran bat spe
and Fig. 4), and recorded the echolocation calls of sixteen (Table 2). We foun
correlations between morphology and echolocation call characteristics. Call
(both highest and lowest frequencies) were correlated with wingloading (r=- 0-62,
P < 0-05 and r= -0-72, P < 0.01 for highest and lowest frequencies, respectively), aspect
ratio (r= -0-54, P< 0.05 and r= -0'59, P< 0.05) and wingtip shape (r=0-51, P <0.05
and r = 0.6, P< 0-05; Fig. 5). Call shape was significantly correlated with wingloading
(r =-0.70, P< 0-05), aspect ratio (r =-0.70, P < 0-05) and wingtip shape (r=0-88,
P < 0-01; Fig. 5). These data indicate that bats with high wingloadings, high aspect ratios
and pointed wingtips normally have echolocation calls dominated by a shallow frequency
modulated (FM) sweep of relatively low frequency, while species with low loadings, low
40, - - T , I 40
1- 16-
E o?o -
8- ? -8
I~' 0
0 0
, 4- -4
._c
2- -2
4 8 16 34 65 100
Mass (g)
FIG. 6. Double log plots showing the relationship betw

line, Y= 1.60X'34) and aspect ratio (open circles and
The circled points represent H. commerso
aspect ratios and rounded wingtips tend to hav

short and broad-band calls. The rhinolophids and hipposiderids belong to the latter
group, but have calls dominated by a constant frequency component. These bats use
Doppler-shifted echoes to detect fluttering prey within clutter, and therefore (in terms of
clutter resistance) their calls are functionally equivalent to the broad-band calls of the
other species.
On the basis of the morphological and echolocation data on the species in this
assemblage we can make predictions about the animals' flight performance and habitat
use. We recognize four broad groups (Fig. 4).
Group 1. Bats with low mass ( < 15 g), low wingloading (6 N m-2 8 N m2) and low
aspect ratio (5-5-6-5), relatively low flight speeds (Vmp (minimum power speed) = 3.5
ms- - 4-5 ms- 1, calculated using eq (10) in Pennycuick 1975), high manoeuvrability and
clutter-resistant echolocation calls. All these species should be manoeuvrable and be able
to forage in cluttered habitats.
Group 2. Bats weighing between 15 g and 25 g, having higher wingloadings (8 N m2 -
12 N m-2) and aspect ratios (6-8), and higher flight speeds (Vmp=4'5 m s-l-5 0 m s-1)
and lower manoeuvrabilities than species in group 1. Rhinolophus hildebrandti uses high
intensity calls dominated by CF components and terminating with a short FM sweep. In
common with other Rhinolophus spp., this bat should be capable of detecting fluttering
prey against background clutter, and although it will be less manoeuvrable than the bats
of group 1, we would expect it to fly in more cluttered habitats than the other species in
group 2. Scotophilus borbonicus and S. dingani use high intensity, 5-10 ms calls with broad
band-width (Table 2). Their calls have two parts, an initial steep FM sweep of c. 40 kHz
followed by a longer shallow terminal FM section. This type of call should enable S.
borbonicus and S. dingani to detect prey both in moderate clutter (the initial portion) and
over relatively long distances (the terminal portion). These species should be relatively
manoeuvrable and we predict they will forage both in open areas and within woodland,
but not within thick vegetation.
Group 3. Only Hipposideros commersoni fits in this group, mainly because of its
TABLE 3. The minimum negotiable distances (MND) for seventeen sympatric
insectivorous bat species resident in the Pafuri region. The minimum inter-string
distance that at least one individual of each species was able to negotiate is also
shown
MND Minimum
(m) (m)
Species n x S.D.
R. hildebrandti 9 0.28 0.07 0.22
R.fumigatus 6 0.35 0.07 0.22
R. landeri 5 0.16 0-04 0-11
R. swinnyi 2 0.19 0.15
H. caffer 13 0.15 0.03 0-11
H. commersoni 2 0.44 - 044
N. thebaica 5 0.12 0.02 0.11
M. bocagei 4 0.28 0.06 0'22
P. rueppellii 5 0.24 0-06 0.15
P. nanus 6 0.20 0.07 0.22
P. rusticus 5 0.24 0.06 0.22
E. capensis 9 0.27 0.09 0.15
E. zuluensis 2 0.30 - 030
N. schlieffeni 9 0.13 0.02 0.11
S. borbonicus 7 0.31 0.10 0.22
S. dingani 6 0.39 0.07 0.30
T. pumila 5 0.44 0.00 0.44
relatively high body mass (c. 90 g, Table 1). The two males we caught were probably
young since typical measurements of mass for this species in Southern Africa are 115-140
g (Smithers 1983). In H. commersoni, wingloading is quite low for its size (Fig. 6) and it has
a lower aspect ratio than we would have predicted (Fig. 6). Nevertheless, it will tend to
have a high flight speed (Vmp = 5 7 m s- ') and relatively low manoeuvrability. The high
mass is somewhat compensated for by the high wing area (= low wingloading), low aspect
ratio and the highly rounded wingtips, all of which should improve manoeuvrability. We
predict that this species will forage in moderate clutter, but will not be capable of entering
thick vegetation. Its echolocation calls are typical of hipposiderids, and assuming that it is
able to compensate for Doppler-shifts, it should be able to detect prey against
background clutter.
Group 4. There are six species in this group: five Tadarida spp. and the emballonurid,
Taphozous mauritianus. They range in mass from 13 g (T. pumila) to 44 g (T. midas), but
they all have high wingloadings (13 N m-2 - 20 N m-2), high aspect ratios (9-15) and
pointed wingtips. This morphological combination will produce high flight speeds
(Vmp=4 8-1 - 5-5 m s-1) and low manoeuvrabilities. These bats use high intensity,
shallow FM calls of low frequency and long duration (c. 15 ms) (Table 2). The association
of high wingloading, high aspect ratio and pointed wingtips with shallow sweep FM calls
suggests that these species should forage in the open where manoeuvrability and prey
detection in clutter are unnecessary.
Manoeuvrability and habitat use

Our results on the manoeuvrability of seventeen species, and the habitat use by nine
species, respectively, support our predictions (Tables 3, 4). There are significant
differences in the bats' abilities to fly through the obstacle courses and in their use of
available foraging space (Friedman's ANOVA X2=33-6, P<0.05 and X2=82, P<0-01,
TABLE 4. Habitat use by nine insectivorous bat species. The percentage time spent in
each zone for each species and the mean habitat use indices are shown. n is the
number of bats released with light-tags, n/o the number of separate observations
and T the total time (s) for which a species was observed. The 'Flight' column
summarizes the species' foraging behaviour. 'Co.' = continuous flight while
foraging and 'Pe.' = short flights from a perch, a number of attacks made during
each flight. The habitat zones are described in Fig. 3. No bats foraged in zone 2
Habitat zones HU
Species n n/, T (1) (3) (4) (5) (6) (7) X S.D. Flight
T. condylura 13 4 114 100 - 1.0 0.0 Co.

T. pumila 12 24 2463 100 - 1.0 0-0 Co.
S. borbonicus 7 6 165 27 - 54-0 18-0 2-0 1-0 Co.
S. dingani 9 12 403 - - 78-0 22-0 4-3 0-6 Co.
H. caffer 20 14 4779 - - 0-3 2-1 3-2 94-4 6-2 0-7 Pe.
R. hildebrandti 7 7 153 62-1 37-9 5.5 0-5 Pe./Co.
E. capensis 20 31 459 74-3 - 22-9 2-8 2-5 1-5 Co.
N. schlieffeni 20 15 228 50-3 - 10-1 38-7 09 2-3 1-0 Co.
P. nanus 0 20 422 13-0 2-0 74-9 9.9 3-5 1-1 Co.
120
(a)
80C
40L
20 Ib) --.,
IE
20 . ( b )~~~~~~~
0-
180 -
Ic)
140
-0
100-
U-
I ~ I ,
60-
20-
0 0-1 0-2 0-3 0-4 0-5 0-6

Minimum negotiable distance (m)
FIG. 7. The relationship between morphology, echolocation call frequency and minimum
negotiable distance. (a) mass (n = 17, Y= -0-06X?27), (b) wingloading (n = 17, Y= - 1-5X?95)
and (c) frequency (n= 13, Y= -0.99X-0?87).
TABLE 5. The insects consumed by twelve sympatric insectivorous bat species

resident in the Pafuri region. n=number of bats; n/i= number of insects; Col.,
Coleoptera; Lep., Lepidoptera; Dip., Diptera; Hem., Hemiptera; Hym., Hymenop-
tera; Tri., Trichoptera; Iso., Isoptera
(a) Taxon
Species n ni Col. Lep. Dip. Hem. Hym. Tri. Iso. 0th.

R. hildebrandti 4 9 33 56 11
R. fumigatus 1 6 53 15 33
H. caffer 3 14 29 21 50
H. commersoni 2 12 100
P. rueppellii 1 1 100
P. rusticus 2 12 67 8 8
E. capensis 5 27 56 22 4 19
E. zuluensis 1 5 40 20 40
E. melckorum 1 4 75 25
N. schlieffeni 3 8 50 13 25 13
S. dingani 4 48 90 2 2 2 2 2 2
T. pumila 4 16 31 19 6 31 13
(b) Size
Species n n/i Large Medium Small
R. hildebrandti 4 9 33 67
R. fumigatus 1 6 33 67
H. caffer 3 14 100
H. commersoni 2 12 100
P. rueppellii 1 1 100
P. rusticus 2 12 100
E. capensis 4 10 20 80
E. zuluensis 1 5 100
E. melckorum 1 4 100
N. schlieffeni 3 8 100
S. dingani 4 48 8 76 16
T. pumila 4 16 100
respectively). On the basis of their mean HU values (Table 4) the bats fell into four
categories:
(a) T. condylura and T. pumila used only zone 1.
(b) N. schlieffeni, E. capensis and P. nanus used a range of habitats, but did not forage
within clutter.
(c) S. borbonicus and S. dingani foraged in zones 4 and 5.
(d) H. caffer and R. hildebrandti foraged in moderate clutter, with H. caffer foraging in
more cluttered habitats. When foraging, both bats sometimes used a perch, from
which they made short flights.
The differences in manoeuvrability are related both to morphology and echolocation
call design. There was a significant correlation between manoeuvrability and habitat use
(rs=0-89, P<0-01), suggesting a causal relationship. Manoeuvrability was negatively
correlated with mass (r= -057, P<0-05) and wingloading (r= -065, P<0-01), and
positively correlated with echolocation call frequency (CF frequency in rhinolophid and
hipposiderid bats, the highest frequency in FM bats; r =0-7, P <001; Fig. 7).
Diet
Our analysis of the diets of twelve species (Table 5) shows no significant interspecific
differences in the proportions of the different insect orders taken by all the bats
(Friedman's ANOVA X2= 4'5, P> 0-05). The diets of most species consisted of moths and
TABLE 6. X2 values for pair-wise comparisons of the diets of six sympatric bat
species
T. pumila N. schlieffeni E. capensis S. dingani H. caffer R. hildebrandti

T. pumila 7.3N.S. 16-3*** 29-3*** 14-8** 8-0
N. schleiffeni 17-6*** 25.4*** 9-7** 10.0
E. capensis 12.7** 14-5*** 12-7***
S. dingani 31.8*** 31-6***
H. caffer 7-9
** Significant at
*** Significant a
N.S., Not signific
beetles in appr
taken in larger
numbers. H. c
generalization,
Trichoptera an
each of the cat
differences in
difference in t
hildebrandti (X2= 7-85, P>0.2, d.f. = 3), but the sizes of beetles found in their faeces
differed significantly (X2= 5.18, P <005, d.f. = 1). E. capensis and N. schlieffeni are
significantly different in diet species composition (X2 = 17-6, P < 0.01, d.f. = 5), but not in
beetle size composition (X2= 1-82, P> 0-05, d.f. = 1).
Although there was no significant difference in the number of insects of different size-
classes taken by all the bats (Friedman's ANOVA X2 = 2-8, P > 0-05), there was a significant
correlation between the size of bat and mean prey size (rs=0-7, P< 0-05). Small bats
tended to take only small prey, but larger bats took prey of a range of sizes (Table 5b).
DISCUSSION
Our results indicate that there is a tendency for differential use of the av
space, with four habitat-use strategies: (i) 'open' foragers (e.g. T. con
pumila); (ii) 'Woodland edge' foragers (e.g. N. schlieffeni, E. capensis an
'Intermediate clutter' foragers (e.g. S. borbonicus and S. dingani); (iv) 'clutt
(e.g. H. caffer and R. hildebrandti). Published data on the foraging b
commersoni (Vaughan 1977) and S. borbonicus (= leucocaster; Barclay 1985)
predictions, while data on the foraging behaviour of H. caffer (Bell
indicate that this species is more versatile than our results would suggest. These
differences in habitat use correlate with differences in the bats' manoeuvrabilities, which
in turn correlates with wing morphology. These data support the first of our hypotheses,
namely that differences in morphology appear to lead to a spatial partitioning of the
available habitat.
A bat's wing morphology will place energetic and mechanical limitations on where it
can forage efficiently. We would expect natural selection to lead to the minimization of
foraging time and energy expenditure while ensuring a maximum rate of energy intake
(R. A. Norberg 1977). The rate of energy intake will be proportional to the number of
insects the bat encounters while foraging, which is likely to increase with increasing flight
speed (R. A. Norberg 1981). One consequence of this is that bats that tend to have higher
flight speeds (high mass, wingloading and aspect ratio) may be able to maintain the same
rate of energy intake in habitats with lower insect densities than bats with lower flight
speeds (low mass, wingloading and aspect ratio). Therefore, bats such as T. condylura and
T. pumila can forage in the open where insect densities tend to be low, while to maintain a
positive energy budget and minimize foraging time, H. caffer has to fly in habitats which
support high insect densities, which in general means flying in highly cluttered habitats
(Lewis 1970; Neuweiler 1983). In general, our data support the suggestion that the lower a
bat's mass, wingloading and aspect ratio the more frequent will be its use of cluttered
habitats.
Species (e.g. H. caffer and N. schlieffeni) that appear to be adapted primarily for
manoeuvrable flight in clutter, may be able to reduce foraging flight costs, without
increasing foraging time and decreasing insect encounter rates, by flying in insect
concentrations that occasionally form outside cluttered areas, for example during the
emergence of the alates of Hymenoptera, Isoptera, Ephemeroptera and Trichoptera, and
on the leeward side of stands of trees where aerial insects tend to collect (Lewis 1970).
Clearly, natural selection will favour those individuals that are behaviourally flexible
enough to make use of these unpredictable situations and, therefore, it should not surprise
us that bats, such as N. schlieffeni and H. caffer, are opportunistic and are not restricted to
foraging in the particular habitats to which they seem most adapted.
Our results confirm the hypothesis that wing morphology and echolocation call design
are parts of the same adaptive complex, and we can predict from this that bats with high
body mass, wingloading and aspect ratio will tend to have low frequency and shallow
band-width calls, which although susceptible to interference from clutter, are ideally
suited for the long-range detection of prey in the relatively open habitats to which these
species are morphologically restricted. On the other hand bats that, for energetic reasons,
are restricted to flying in habitats of high insect density, and therefore normally high
clutter, use broad band-width (or constant frequency calls plus an ability to compensate
for Doppler-shifts) calls of relatively high frequency which although only effective over
short range are clutter-resistant.
It is clear that the characteristics of a particular call are very important in determining a
bat's ability to detect and locate its prey in a particular habitat, and if a particular bat
species was capable of generating only one type of call, and interpreting that call in only
one way, it would be restricted to hunting in a particular habitat. However, there is a great
deal of evidence to suggest that bats are very flexible in their use of echolocation
(Simmons et al. 1978; Fieldler 1979; Habersetzer 1981; Bell 1982, 1985; Neuweiler 1983),
and can therefore circumvent the restrictions imposed by a particular combination of
echolocation call characteristics. We therefore propose that the primary mechanism of
resource partitioning in bat communities will be wing morphology and therefore
manoeuvrability, with echolocation behaviour being varied to suit a particular foraging
situation.
Our data on the diets of these bats support the hypothesis that bats with similar
strategies of habitat use tend to have similar diets and vice versa (Hespenheide 1975).
While our data on prey size support the general ecological principle that the larger the
predator the larger the prey is likely to be, they also show that the larger species tend to
take prey from a wider range of sizes, presumably because they can easily subdue small
insects, but that the reverse is not true. These data support the idea that the primary
mechanism of resource partitioning in this community at this time of year is flight
performance, with competition for food being weak or absent. It is possible, however, that
those bats with the same habitat use strategies and the same diets show differences in flight
periodicity. The majority of species were caught within the first 2 h after sunset, suggesting
that flight times were similar, although we did notice that some species were always caught
earlier than others. The fact that R. hildebrandti, S. borbonicus and T. midas tended to
forage at the same time suggest that, in general, foraging times substantially overlap
(Fenton & Rautenbach 1987). Moreover, there is little evidence of temporal partitioning
of foraging time in sympatric insectivorous bats (Fenton 1982, 1985).
The data presented here thus suggest that differences in wing morphology and
echolocation call characteristics lead to spatial resource partitioning in insectivorous
bats, and that measurements of morphology and echolocation call characteristics are
good predictors of the ecological structure of a community.
ACKNOWLEDGMENTS
We would like to thank the following people for their help in the field:
Balcombe, Dr L. E. 0. Braak, R. M. Brigham, Dr M. B. Fenton, L. A. Hare, Dr G. L.
Holroyd, Dr J. Ostwald, Dr H.-U. Schnitzler, M. G. Stoneman and V. Wai-Ping. Drs
M. B. Fenton, U. M. Norberg, T. H. Kunz, T. H. Fleming and two anonymous referees
read earlier drafts of this paper and we are grateful for their comments. We also thank Dr
R. Thomas for his statistical advice. Finally we would particularly like to thank the
Warden of the Kruger National Park, Dr U. de V. Pienaar and Dr V. de Vos (Assistant
Head: Research), for permission to conduct this research in the Park and for the use of the
facilities at Pafuri. H. D. J. N. A. is a Science and Engineering Research Council (U.K.)
Research Fellow. This work was financed from a grant awarded to Dr M. B. Fenton by
the Natural Sciences and Engineering Research Council of Canada and from funds
awarded to I. L. R. by the Foundation of Research Development of the Council for
Scientific and Industrial Research, Pretoria.
REFERENCES
Aldridge, H. D. J. N. (1986a). Manoeuvrability and ecology in British bats. Myotis, 23-24, 5

Aldridge, H. D. J. N. (1986b). Manoeuvrability and ecological segregation in the little brown
and Yuma (M. yumanensis) bats. Canadian Journal of Zoology, 64, 1878-1882.
Barclay, R. M. R. (1985). Foraging behaviour of the African insectivorous bat, Scotoph
Biotropica, 17, 65-70.
Bell, G. P. (1982). Behavioral and ecological aspects of gleaning by a desert insectivorous bat
(Chiroptera: Vespertillionidae). Behavioural Ecology and Sociobiology, 10, 217-223.
Bell, G. P. (1985). The sensory basis of prey location by the California leaf-nosed bat Macr
(Chiroptera: Vespertilionidae). Behavioral Ecology and Sociobiology, 16, 343-347.
Bell, G. P. & Fenton, M. B. (1984). The use of Doppler-shifted echoes as a flutter detection and c
system: the echolocation and feeding behavior of Hipposideros ruber (Chiroptera:
Behavioural Ecology and Sociobiology, 15, 109-114.
Buchler, E. R. (1976). A chemiluminescent tag for tracking bats and other small nocturnal
Mammalogy, 57, 173-176.
Farney, J. & Fleharty, E. D. (1969). Aspect ratio, loading, wing span and membrane areas of
Mammalogy, 50, 362-367.
Fenton, M. B. (1982). Echolocation, insect hearing, and feeding ecology of insectivorous bats. Ecology of Bats
(Ed. by T. H. Kunz), pp. 261-280. Plenum Press, New York.
Fenton, M. B. (1985). The feeding behaviour of insectivorous bats: echolocation, foraging strategies and
resource partitioning. Transvaal Museum Bulletin, 21, 5-16.
Fenton, M. B. & Rautenbach, I. L. (1987). The roosting and foraging behaviour of three species of African
insectivorous bats (Rhinolophus hildebrandti: Rhinolophidae; Scotophilus borbonicus: Vespertilionidae; and
Tadarida midas: Molossidae). Canadian Journal of Zoology, 64, 2860-2867.
Fiedler, J. (1979). Prey catching with and without echolocation in the Indian False Vampire bat (Megaderma
lyra). Behavioral Ecology and Sociobiology, 6, 155-160.
Findley, J. S., Studier, E. H. & Wilson, D. E. (1972). Morphologic properties of bat wings. Journal of
Mammalogy, 53, 429-444.
Findley, J. S. & Wilson, D. E. (1982). Ecological significance of Chiropteran morphology. Ecology of Bats (Ed.
by T. H. Kunz), pp. 243-260. Plenum Press, New York.
Gaisler, J. (1959). Beitrag zum vergleichenden studium des Flugapparates der Fledermause (Microchiroptera).
Folia Zoologica, 8, 37-62.
Griffin, D. R. (1971). The importance of atmospheric attenuation for the echolocation of bats (Chiroptera).
Animal Behaviour, 19, 55-61.
Habersetzer, J. (1981). Adaptive echolocation sounds in the bat Rhinopoma hardwickei, a field study. Journal of
Comparative Physiology, 144, 559-566.
Hesphenheide, H. A. (1975). Prey characteristics and predator niche width. Ecology and Evolution of
Communities (Ed. by M. L. Cody & J. M. Diamond), pp. 158-180. Belknap Press, Cambridge,
Massachusetts.
Lewis, T. (1970). Patterns of distribution of insects near a windbreak of tall trees. Annals of Applied Biology, 65,
213-220.
Neuweiler, G. (1983). Echolocation and adaptivity to ecological constraints. Neuroethology and Behavioural
Physiology. Roots and Growing Points (Ed. by F. Huber & H. Markl), pp. 280-302. Springer-Verlag, Ber
Norberg, R. A. (1977). An ecological theory on foraging time and energetics and choice of optimal
searching method. Journal of Animal Ecology, 46, 511-529.
Norberg, R. A. (1981). Optimal flight speeds in birds when feeding young. Journal of Animal Ecology, 50,
477.
Norberg, U. M. (1981). Allometry of bat wings and legs and comparison with bird wings. Philosophical
Transactions of the Royal Society (London), B, 287, 131-165.
O' Shea, T. J. & Vaughan, T. A. (1980). Ecological observations on an east African bat community. Mammalia,
44, 485-496.
Puranik, R. G., Gopalakrishna, G. & Chari, N. (1976). Flight parameters of the short-nosed fruit bat Cynopterus
marginatus. Proceedings of the Indian Academy of Sciences, B, 83, 160-165.
Pennycuick, C. J. (1975). Mechanics of flight. Avian Biology. Vol. 5 (Ed. by D. S. Farner, J. R. King & K. C.
Parkes), pp. 1-73. Academic Press, New York and London.
Simmons, J. A., Fenton, M. B., Ferguson, W. R., Jutting, M. & Palin, J. (1979). Apparatus for research on animal
ultrasonic signals. Life Sciences Miscellaneous Publications. Royal Ontario Museum, Toronto. pp. 1-31.
Simmons, J. A., Lavender, W. A., Lavender, B. A., Childs, J. E., Hulebak, K., Rigden, M. R., Sherman, J.,
Woolman, B. & O'Farrell, M. J. (1978). Echolocation by free-tailed bats (Tadarida). Journal of Comparative
Physiology, 125, 291-299.
Simmons, J. A. & Stein, R. A. (1980). Acoustic imaging in bat sonar: Echolocation signals and the evolution of
echolocation. Journal of Comparative Physiology, 135, 61-84.
Smith, J. D. & Starrett, A. (1979). Morphometric analysis of chiropteran wings. Biology of bats of the New World
family Phyllostomatidae (Ed. by R. J. Baker, J. K. Jones, Jr & D. C. Carter), pp. 1-441. Special Publications
of the Museum of the Texas Technical University, 16. Lubbock.
Smithers, R. H. N. (1983). The Mammals of the Southern African Subregion. University of Pretoria, Pretoria.
Tuttle, M. D. (1974). An improved trap for bats. Journal of Mammalogy, 55, 475-477.
Vaughan, T. A. (1959). Functional morphology of three bats: Eumops, Myotis, Macrotus. Publications of the
University of Kansas Museum of Natural History, 12, 1-153.
Vaughan, T. A. (1966). Morphology and flight characteristics of molossid bats. Journal of Mammalogy, 47, 249-
260.
Vaughan, T. A. (1977). Foraging behaviour of the giant leaf-nosed bat (Hipposideros commersoni). East African
Wildlife Journal, 15, 237-249.
(Received 18 June 1986)

Aldridge y Rautenbach 1987 Morphology Ecolocacion and Food Partitioning in Insectivorous Bats

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Morphology, Echolocation and Resource Partitioning in Insectivorous Bats

Author(s): H. D. J. N. Aldridge and I. L. Rautenbach

MORPHOLOGY, ECHOLOCATION AND RESOURCE

BY H. D. J. N. ALDRIDGE* AND I. L. RAUTENBACH

Department of Biology, Carleton University, Ottawa KIS 5B6, Canada

The study of resource partitioning in bat communities has concentrated primarily on

MATERIALS AND METHODS

We caught twenty-six species of the sub-order Microchiroptera either in mist-nets or in

TABLE 1. Morphological parameters of twenty-six sympatric insectivorous bat

Mass Wingloading Aspect ratio Wingtip

balance (accurate to 0-1 g; Ohau

The fundamental measurements were used to derive three further parameters:

TABLE 2. Characteristics of the search-phase echolocation calls of sixteen sympatric

(Buchler 1976). After the light-tags were attached t

wherefis change in call frequen

80- 160-- -24

FIG. 5. Double log plots showing the

Manoeuvrability and habitat-use predictions

FIG. 6. Double log plots showing the relationship betw

aspect ratios and rounded wingtips tend to hav

Manoeuvrability and habitat use

T. condylura 13 4 114 100 - 1.0 0.0 Co.

0 0-1 0-2 0-3 0-4 0-5 0-6

TABLE 5. The insects consumed by twelve sympatric insectivorous bat species

Species n ni Col. Lep. Dip. Hem. Hym. Tri. Iso. 0th.

T. pumila N. schlieffeni E. capensis S. dingani H. caffer R. hildebrandti

Aldridge, H. D. J. N. (1986a). Manoeuvrability and ecology in British bats. Myotis, 23-24, 5

(Received 18 June 1986)

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