PREDATION UPON MOTHS BY FREE-FORAGING
HIPPOSIDEROS CAFFER
D.
C.
DUNNING AND MARTIN KROGER
Department of Biology, West Virginia University, P.O. Box 6057 Morgantown,
WV 26506-6057 (DCD)
Transvaal Museum, P.O. Box 413 Pretoria, South Africa (MK)
Predation upon insects by the bat Hipposideros caffer was studied at Skukuza in the Kruger
National Park of South Africa. These bats, whose echolocation calls are inaudible to moths,
fed overwhelmingly upon tympanate Lepidoptera, although insects of other orders frequently dominated the local light-responsive, flying, nocturnal insect community. The relative numbers of noctuid, pyralid, and arctiid moths taken by the bats were proportional
to the representation of these families in the general population of moths, but they took
disproportionately fewer moths of the family Geometridae. The bats also ate significantly
fewer arctiid moths of those species capable of clicking than of those species that could
not. Because the arctiids could not hear approaching H. caffer, these moths did not click
before contact with an attacking bat, and their clicks could not protect them by interfering
with echolocation by the predators. These results are consistent with the startle and acousticaposematism hypotheses for the bat-protective function of arctiid clicks.
Key words:
Hipposideros caffer, acoustic defenses, predation preferences, echolocation
Insectivorous bats usually prey upon a
variety of insects (Fenton, 1985), but some
species may specialize on insects of one or
a few orders, particularly in the presence of
competitors (Findley, 1993). Tympanate
moths and several other night-flying insects
can hear the echolocation pulses of bats and
take evasive action, which makes them difficult to capture (Miller, 1975; Moiseff and
Hoy, 1983; Robert et aI., 1992; Roeder and
Treat, 1961; Yager and Hoy, 1986; Yager et
aI., 1990). In tum, many bats feeding upon
insects that can hear ultrasonic pulses have
modified their echolocation behavior so
their prey cannot detect them (Fenton,
1982). Hipposideros caffer, a moth-eating
bat (Bell and Fenton, 1984; Findley and
Black, 1983) from southern Africa, uses
narrow bandwidth pulses at 140 kHz, well
above the known hearing range of any moth
(Fenton, 1986; Fenton and Fullard, 1979;
Fullard, 1987). Prey taken by this bat may
indirectly reveal the consequences of acoustic interactions between the predator and
Journal of Mammalogy, 77(3):708-715, 1996
various families of tympanate moths, especially the Arctiidae.
Some moths of the family Arctiidae, including the Ctenuchinae (ScobIe, 1992),
produce trains of ultrasonic clicks using
metepisternal tymbal organs, in response to
tactile stimulation and when they hear approaching bats (Blest, 1964; Blest et aI.,
1963; Dunning, 1968; Fullard, 1984).
Moths of this family are less likely than
other tympanate moths to take evasive action in response to bat-like ultrasonic stimulation (Dunning et al., 1992). Both captive
and free-foraging bats eat fewer arctiids
than are available to them (Acharya and
Fenton, 1992; Dunning, 1968; Dunning and
Kriiger, 1995; Dunning et al., 1992). Foraging red bats usually do not catch moths
that can click, although they may catch and
then drop those of the same species that
have been experimentally muted (Acharya
and Fenton, 1992; Dunning et aI., 1992). It
appears that many arctiids are distasteful or
malodorous to bats and advertise their unpalatability with their clicks (i.e., these
708
August 1996
DUNNING AND KRUGER-PREDATION ON MOTHS BY HIPPOSIDEROS
moths are acoustically aposematic-Dunning, 1968; Dunning et aI., 1992; Surlykke
and Miller, 1985). In addition, the clicks
may startle (Bates and Fenton, 1990; Stoneman and Fenton, 1988) or acoustically confuse (Fullard et aI., 1979; Miller, 1991; Surlykke and Miller, 1985) attacking bats, by
masking echoes or by jamming echolocation capabilities. These three hypotheses
(acoustic aposematism, startle, and acoustic
confusion) are not mutually exclusive and
may operate simultaneously or in different
interactions of bats and moths.
Arctiid moths that cannot hear attacking
bats will not click, at least not until they are
touched. Because the acoustic-confusion
hypothesis requires that clicks of moths be
emitted before contact, when they might interfere with the attacking bat's abilities to
find and catch the insects, H. caffer should
not be deterred from capturing and eating
palatable arctiids, whether or not the moths.
have tymbal organs (Fullard et aI., 1994).
Therefore, if clicks operate only by interfering with echolocation of attacking bats,
we would expect H. caffer to eat about the
same proportion of arctiid moths as are
present in the general popUlation of moths
regardless of the clicking abilities of the
moths.
The sudden emission of clicks by a moth
upon contact with some portion of the
catching surface (Webster and Griffin,
1962) of an attacking bat may startle H. caffer enough to effect the moth's escape. Protection against bats by this mechanism
should operate only if the clicking moths
are relatively rare members of the lepidopteran population (Endler, 1991). At larger
proportions, any protection that clicking
moths derive may be due to acoustic aposematism. In model experiments using
trained bats feeding upon palatable prey,
Eptesicus fuscus was seldom startled by
clicks of moths after two or three trials
(Bates and Fenton, 1990; Miller, 1991).
However when similar experiments were
performed with the gleaning bat Macrotus
californicus, predators consistently were
709
deterred by clicks of moths, even after
many trials (Stoneman and Fenton, 1988).
Because H. caffer ecologically is more similar to Macrotus than to Eptesicus in that
both are gleaners, it may be that these Hipposideros would not habituate to clicks of
moths and would be startled even if clicking arctiids were relatively common. Therefore, both the startle and acoustic-aposematism hypotheses may explain a preference for non-clicking arctiids by H. caffer,
regardless of the relative abundance of
these moths.
MATERIALS AND METHODS
A group of H. caffer that feeds on insects in
the vicinity of the Nature Conservation Centre
at Skukuza in the Kruger National Park of South
Africa were observed flying into an attic area
between the ceiling and the roof of the Centre,
where they dropped such inedible fragments as
the wings of their prey. Because the wings of
moths could be used to determine their identity,
insects taken by these free-foraging H. caffer
were compared with moths and other insects
captured in light traps, to determine feeding
preferences of the bats.
The floor of the attic area was covered with
plastic sheeting at the beginning of the study.
Insect remains not eaten by bats the previous
night and dropped on the floor were collected
from this area on 26 days in November and December of 1993, sorted to order of insect, and
counted. Fragments of lepidopterans (mostly
wings) were identified to family and sometimes
to species, whenever possible, according to Van
and Kroon (1986). All recognizable remains of
moths of the family Arctiidae were identified to
species. Bats apparently did not use any particular part of the area as a night roost, because
both feces and insect remains were distributed
over the entire 50.6-m2 area, rather than concentrated in a few places. Fragments of insects collected from the attic area each day are hereafter
referred to as the samples of insects eaten by
bats.
Identity of bats and the areas they frequented
were established with an Ultrasound Advice
Mini-2 bat detector tuned to 30, 40, and 140
kHz, using a directional cone when necessary to
determine where bats with different echolocation signals were flying. Times of maximum ac-
7lO
JOURNAL OF MAMMALOGY
tivity of H. caffer were established by listening
with the bat detector tuned to 140 kHz at hourly
intervals throughout one night, and activity was
sampled twice each night on 10 additional
nights. All fecal pellets were collected on the 1st
day and examined for the presence of lepidopteran scales. Thereafter, any fresh fecal pellets
collected with the fragments of insects were examined similarly.
Insects were collected during times of maximum activity of H. caffer (2200-0330 h) in a
suspended, ultraviolet light-trap and promptly
killed with cyanide supplemented with chloroform. On the last 2 nights insects were collected
between full dark (1900 h) and 0.5 h before
dawn (0330 h). About 500 m away, the trap was
located in an area similar in height and abundance of vegetation and in abundance of external lights to the vicinity of the building where
the insect remains dropped by the bats were collected.
On most nights, large insects with lengths of
body or forewings >25 mm were removed, sorted to order, and counted. If the remaining insects
were numerous enough to completely cover at
least one-half of a 43.5- by 32.5-cm sorting tray,
their numbers and distributions of orders were
estimated by sorting and counting 12.5-25.5%
of them; otherwise all were sorted and counted.
All Lepidoptera with lengths of forewing >5
mm were identified to family, and all arctiids
and some moths of other families were identified
to species according to Vari and Kroon (1986),
using the lepidopteran collection at the Transvaal Museum. Insects with lengths of forewing
or body <2 mm were not counted, because their
remains were too small to have been detected if
present in the samples of insects eaten by bats.
On 1 night, the light-trap sample was dominated
by a large number of alate termites, and only
12.5% of the whole collection was processed.
Hereafter, insects collected in the light trap and
processed as described are referred to as the
light-trap samples.
Rainfall, relative humidity, wind, and temperatures taken at a weather station about 100m
from the location of the light trap and 300 m
from the Nature Conservation Centre (where the
samples of insects eaten by bats were collected)
were correlated with the overall size of the lighttrap sample and with the proportions of different
orders of insects in it. Proportions of insects of
different orders, of different families of Lepi-
Vol. 77, No.3
doptera, and (on 10 nights) of one species in
each of the two most common families of moths
in the 2 nightly samples (the light trap and the
fragments dropped by bats) were compared using G tests.
Live arctiids of the species in the light trap
and samples of insects eaten by bats were caught
at another light and examined for the presence
of metepistemal tymbal organs. They were then
assayed for production of clicks in response to
tactile stimulation. Proportions of those species
that proved capable of clicking were compared
with those that never clicked (and which lacked
tymbal organs) in the light-trap samples and
samples of insects eaten by bats, using G tests.
RESULTS
Light-trap samples.-A total of 117,022
insects was collected in the light trap during
the 26 nights of the study, almost one-half
of them (57,702) on the only night with a
hard rain. The overall number of insects
(Fig. 1) in the light-trap sample each night
was strongly influenced by rainfall (F =
210.66, d.! = 12, P < 0.001). Most of this
correlation is a consequence of the emergence of enormous numbers of alate termites on the night with a heavy rain (F =
39,411.78, d.! = 8, P < 0.001) and of alate
ants on nights of moderate rainfall (F =
99.19, d.! = 8, P < 0.001). The numbers
of beetles and moths, which usually dominated the catch, were not affected by rainfall episodes (Coleoptera: F = 2.54, d.!
8, P = 0.07; Lepidoptera: F = 0.75, df =
8, P = 0.65).
Samples of insects eaten by bats.-AIthough bats with echolocation signal-frequencies detected at 30-40 kHz were abundant around the building where the fragments dropped by bats were collected and
in adjacent areas, such bats (probably Tadarida pumila) did not approach to within
ca. 3 m of the opening to the attic area. The
only signals from bats detected in and near
the attic area were those characteristic of H.
caffer, at a bat-detector center-frequency of
140 kHz. Bats emitting these cries were observed each evening, flying in a recognizable looping path along the front of the
August 1996
DUNNING AND KROGER-PREDATION ON MOTHS BY H1PPOSlDEROS
711
4 0 . . . . - - - - - - - - - - - - - - - - - - - - - - - - - - - -........ 60,800
........
Rainfall
--0-- Llght.trap sample
50,800
.:s
30
40,800
!-.;:a
r
Col
li
30,800
20
.1
'S
Js
=c
20,800
10
:I
III
iEo-
10,800
o
10
20
Night number
FIG. I.-Effect of rainfall on the number of insects captured in the light trap.
building, into the attic area, and out again.
There were at least six individuals among
these bats, but the total number was not determined.
Few live insects were observed in the attic area where the samples of insects eaten
by bats were collected, most of which were
mud-dauber wasps (Hymenoptera: Sphecidae). Spiders and harvestmen (Chelicerata:
Arachnida: Araneae and Opiliones) also
were apparent residents of the attic area.
A total of 779 fragments of insects was
collected from the attic area in the 26 nights
of the study, 720 (92.4%) of which were
remains of lepidopterans. All fecal pellets
examined contained lepidopteran scales.
Neither the relative abundance of remains
of insects nor the proportion of Lepidoptera
among them reflected the abundance of insects or the proportion of Lepidoptera in the
light-trap samples on the same 26 nights
(Figs. 2 and 3). Proportions of insects in all
15 orders represented in this study were sig-
nificantly different in the two samples (G
= 2,914.05, d.f. = 14, P < 0.001).
Among the 720 fragments of lepidopterans in the samples of insects eaten by bats,
546 were identified at least to family. The
proportion of geometrids in the sample of
insects eaten by bats was significantly less
than the proportion of the same family in the
light-trap samples (G = 14.79, dj. = 1, P
< 0.001), but none of the other three families found in both samples (Arctiidae, G =
0.151; Noctuidae, G = 0.235; and Pyralidae,
G = 0.647) were significantly different.
Families common in the light-trap samples
that were not represented in the samples of
insects eaten by bats included Agaristidae,
Lasiocampidae, Lymantriidae, Notodontidae, and Saturniidae. Only two sphingid
fragments were found in the samples of insects eaten by bats, but 151 moths of the
family Sphingidae were captured in the light
trap. Four wing fragments from moths of the
family Tineidae were found among the re-
712
JOURNAL OF MAMMALOGY
Vol. 77, No.3
l00,oOO-r---------------------------------r7o
· .. 0··· Light.trap sample
.....e--- Samples of insects eaten by
60
bats
so
10,000
1
]
.5
40
·
·
o-.d
1,000
30
20
l00+----------T----------~--------~----------,_--------~--LI0
o
10
20
Night number
FIG. 2.-Numbers of insects in the sample from the light-trap and of insect remains dropped by
foraging Hipposideros caffer on each night of the study. Because of strong effects of rainfall, the
numbers of insects in the sample from the light-trap were highly variable, so the logarithms of these
numbers have been plotted.
l00~----------__~--------------------~~----------------~__------__,
80
•
60
... 0 . ..
40
·
···
o.,
···
··...
···
····· '.q·
o
·
.:.,..... :.. · ...9
():. '8 ·· o..
··· .
.
! ,:
0:
Samples of insects eaten by bats
Light-trap sample
....
20
0.
.0....
···
·
·...
·
· ·0·"a•.
o.
'O•• q
·:··! ~...
·· o. ..
··
·· ·· ···
··..···
·
Q
~
~
!:
o+------u'~c·~·O~·-·--------__r--~>_----~----------~--------~~
o
10
20
Night number
FIG. 3.-Proportion of Lepidoptera in the two samples on each night of the study.
August 1996
DUNNING AND KROGER-PREDATION ON MOTHS BY H1PPOSIDEROS
mains dropped by the bats, but no tineids
comparable in size to those caught by the
bats were captured in the light trap.
Although proportions of moths of three
of the most common families were not significantly different in the light-trap and
samples of insects eaten by bats, within
these families, proportions of some species
or groups of species were different. The
bats caught a significantly larger proportion
of the noctuid Prodotis stolida (G = 60.25,
d.! = 1, P < 0.001) and the pyralid Tyndis
dentilinealis (G = 189.93, d.! = 1, P <
0.001) than were represented in the lighttrap sample. Although arctiids were similarly rare in both samples, bats caught a
smaller proportion (G = 4.23, d.! = 1, P
< 0.05) of those species capable of producing ultrasonic clicks than of those that could
not, although the two groups were about
equally represented in the light-trap sample.
DISCUSSION
Because Hipposideros caffer fed mainly
upon moths whose numbers were not affected by episodes of rainfall, there was no
correlation between rainfall and predation
of H. caffer. Although the foraging range
of H. caffer is unknown, because of its low
aspect ratio, broad wings, and consequent
slow, relatively inefficient, flight (Fenton,
1992), it probably does not forage widely.
The bats were consistently observed to enter the attic area from a direction opposite
to the location of the light trap, suggesting
that they may have captured their moths
>500 m from the light trap. We detected no
signals at 140 kHz in our occasional samples of the bats flying around the light trap.
Because the trap was in an area ecologically
similar to that around the Nature Conservation Centre it appears likely that the insects caught in the light trap and those captured by the bats represented independent
samples of the same general population of
insects.
Although H. caffer is known to feed primarily upon Lepidoptera (Bell and Fenton,
1984), the extent of this specialization has
713
not been documented previously. Because
the light trap sampled only those nocturnal
insects attracted to ultraviolet light, and
moths are attracted more than insects of
some other orders, the light-trap sample
probably was skewed in favor of Lepidoptera. Even so, the proportion of non-lepidopteran insects caught in the light trap always exceeded that in the sample of insects
eaten by bats (Fig. 3). Even on nights when
Lepidoptera were a small proportion of the
insects captured in the light trap, e.g., on
nights 3-5 and on night 12, they comprised
2=75% of the diet of these bats (Fig. 3). H.
caffer evidently could capture insects of other orders, for 59 fragments from non-Iepidopterous insects were collected in the samples of insects eaten by bats. The absence of
remains of mud-dauber wasps, spiders, and
harvestmen in the fragments dropped by the
bats indicates that prey was caught elsewhere and carried into the attic area.
Moths do react to ultraviolet light, so, insofar as the area around the light trap was
ecologically similar to that where the bats
hunted, the sample from the light trap may
have accurately represented the Lepidoptera
available in the general area. If this is so,
these bats did not take a random sample of
the insects available even within the order
Lepidoptera. This also was true of red bats
(Lasiurus borealis) foraging on Lepidoptera
in Canada (Dunning et al., 1992). Like the
Nearctic bats, H. caffer took about the same
proportion of noctuids as were captured in
a light trap, but unlike L. borealis, which
seemed to prefer geometrids, these H. caffer
appeared to discriminate against them.
Hearing ranges of some geometrids are
much higher than those of noctuids (Miller
and Surlykke, in press), so the African geometrids may have been able to hear approaching H. caffer and evade the bats.
The strong preferences of H. caffer for
one species in each of the two most common families of moths has not been observed previously. Although these two species were common in the sample from the
light trap, they were a much larger propor-
714
JOURNAL OF MAMMALOGY
tion of the wings collected in the sample of
insects eaten by bats. The bases of these
preferences are unknown.
In March 1992, a 1.3-m2 sample of the
remains of insects dropped by H. caffer in
the 50.6-m2 attic area included 124 moth
wings, none of which were those of arctiids
(Dunning and KrUger, 1995). The interval
over which these remains had been dropped
was unknown. In the present study, arctiids
comprised 2.2% of all the moths captured
in the light trap and 2.4% of the lepidopteran remains in the sample of insects eaten
by bats. If arctiids were 2-3% of the moths
available during the time that the 1992 collection accumulated, and if H. caffer took
arctiids in proportion to their representation
in the general population of moths, as the
bats seemed to do in this study, we should
have found two or three arctiid wings in the
1.3-m2 sample. Two or three wings easily
could have been missing from a single sample. Therefore the absence of remains of
arctiids in the 1992 study may have been
the result of sampling error.
Arctiids represented a larger proportion
of the moths captured in the light trap in
the Canadian study than in the present one,
but were seldom taken by the foraging red
bats (Dunning et al., 1992). The red bats
did catch (and usually dropped) a larger
proportion of experimentally-muted arctiids
tossed into their paths than of conspecifics
capable of clicking, suggesting that the
clicks of moths prevented contact with the
bats (Acharya and Fenton, 1992; Dunning
et al., 1992). There were relatively small
numbers of arctiids in either of the samples
in the present study, but the proportions
were similar. This suggests that H. caffer
did not discriminate against all arctiids as
did the free-foraging red bats and captive
bats of several species that have been tested
(Dunning, 1968; Dunning and Kruger,
1995; Dunning et aI., 1992), which supports
the acoustic-confusion hypothesis.
Fullard et al. (1994) suggested that clicks
of arctiids do not protect moths against
gleaning bats such as H. caffer, because
Vol. 77, No.3
these insects cannot hear the approaching
bats and produce clicks that might be interpreted as phantom echoes. Comparison of
the proportions of clicking and non-clicking
species of arctiids in the light-trap sample
and among the fragments of arctiids in the
sample of insects eaten by bats showed that
H. caffer preferentially took those without
tymbals, suggesting that an ability to click
did protect the moths. These observations
are not consistent with the acoustic-confusion hypothesis, although the number of remains of arctiids in the sample of insects
eaten by bats was too small to completely
rule out this hypothesis.
Because arctiids were uncommon among
the moths caught in this study, it may be
that exposure to bats was rare enough that
the startle effect of these deimatic sounds
(Edmunds, 1974) would have been sufficient to protect the moths (Endler, 1991).
However, previous experiments with captive T. pumila in the same area (Dunning
and KrUger, 1995) suggested that some of
these moths may have been unpalatable and
thus protected from predation, and these
data do not rule out the acoustic-aposematism hypothesis. Thus, the results of this
study are consistent with both the acousticaposematism and the startle hypotheses for
the bat-protection function of clicks of arctiids, but inconsistent with the acoustic-confusion hypothesis.
ACKNOWLEDGMENTS
We thank the National Parks Board of the Republic of South Africa for permission and partial
support of this project. We also thank L. E. O.
Braack, Senior Research Officer of the Kruger
National Park, and staff of the Nature Conservation Centre at Skukuza. I. L. Rautenbach, Director of the Transvaal Museum, was helpful in
obtaining permission for this study. C. Bennington and W. V. Thayne of West Virginia University helped with the statistical analysis. We thank
K. KrUger for hospitality to D. C. Dunning in
Pretoria, K. Garbutt for permitting her 2 weeks
of leave from West Virginia University, and R.
P. Sutter for teaching one of her classes during
field work.
August 1996
DUNNING AND KROGER-PREDATION ON MOTHS BY HIPPOSIDEROS
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Submitted 12 December 1994. Accepted 31 October
1995.
Associate Editor was Karen McBee.