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Variation in the echolocation calls of the hoary bat (Lasiurus cinereus): Influence
of body size, habitat structure, and geographic location

Article  in  Canadian Journal of Zoology · September 1999

DOI: 10.1139/cjz-77-4-530

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530

Variation in the echolocation calls of the hoary

bat (Lasiurus cinereus): influence of body size,
habitat structure, and geographic location
Robert M.R. Barclay, James H. Fullard, and David S. Jacobs

Abstract: The echolocation calls of bats vary according to the task being undertaken by the individual. Within species,
there is evidence for geographic, habitat, and individual variation, although it is often difficult to separate these factors
and rule out variation in recording or analysis techniques. We studied the variation in echolocation call design of the
hoary bat (Lasiurus cinereus) by analyzing calls, recorded in the same manner, from free-flying bats at one site in
Manitoba and at four sites in the Hawaiian islands. Sites varied in terms of the proximity of the bats to vegetation. As
predicted, individuals from the larger subspecies (Lasiurus cinereus cinereus; Manitoba) used lower frequency calls
than did the smaller individuals in Hawai′i. Within the same habitat type (open or closed), there was evidence for
differences in echolocation call design between populations on different islands. On the island of Hawai′i, bats at a
single site used shorter higher frequency calls when foraging within the vegetation than when foraging in the open,
again as predicted. However, bats foraging in different sites used calls with the opposite characteristics to those
predicted on the basis of the openness of the site. Although there are several possible explanations for this, we suggest
that prey encounter rate and the ability of bats to augment acoustic prey detection with visual information may be
important.
Résumé : Les cris d’écholocation des chauves-souris varient selon leur fonction. Au sein d’une espèce, la variation est
tributaire du lieu géographique, de l’habitat et de l’individu, bien qu’il soit difficile de distinguer ces facteurs et
impossible de rejeter l’hypothèse d’une variation générée par les techniques d’enregistrement ou d’analyse. Nous avons
étudié la variation dans le système des cris d’écholocation de la Chauve-souris cendrée (Lasiurus cinereus) par analyse
de cris enregistrés de la même façon chez des chauves-souris libres à un site au Manitoba et à quatre sites dans les
îles Hawai′i. Les différences entre les sites résidaient surtout dans la distance séparant les chauves-souris de la
végétation. Tel que prédit, les individus de la plus grosse sous-espèce (Lasiurus cinereus cinereus, Manitoba) utilisaient
des cris de fréquence plus basse que les individus plus petits d’Hawai′i. Dans un même type d’habitat (ouvert ou
fermé), les systèmes de cris d’écholocation variaient chez les populations de différentes îles. Sur l’île d’Hawai′i, les
chauves-souris à un site utilisaient des cris plus courts et de plus haute fréquence lorsqu’ils chassaient dans la
végétation que lorsqu’ils chassaient en terrain ouvert, tel que prédit. Cependant, les chauves-souris chassant à d’autres
sites utilisaient des cris aux caractéristiques opposées à celles prévues dans un habitat ouvert. Bien qu’il y ait plusieurs
explications possibles à ce paradoxe, nous croyons que la fréquence des rencontres avec des proies et la capacité des
chauves-souris e compléter leur système de détection acoustique par des signaux visuels sont sans doute des facteurs
importants.

[Traduit par la Rédaction] Barclay et al. 534

Introduction of different species (Aldridge and Rautenbach 1987; Nor-

Microchiropteran bats use a variety of echolocation call
designs. Among aerial insectivorous species, echolocation
call variation is related to the body size and foraging habitat
Received May 25, 1998. Accepted December 18, 1998.
R.M.R. Barclay.1 Department of Biological Sciences,
University of Calgary, Calgary, AB T2N 1N4, Canada.
J.H. Fullard. Department of Biology, Erindale College,
University of Toronto, Mississauga, ON L5L 1C6, Canada.
D.S. Jacobs.2 Department of Zoology, University of Hawai′i
at Manoa, 2538 The Mall, Honolulu, HI 96822, U.S.A.
1 of targets and are better for detecting small prey (Simmons
Author to whom all correspondence should be addressed
(e-mail: barclay@ucalgary.ca).
2
Present address: Department of Zoology, University of Cape
Town, Private Bag, Rondebosch 7700, Cape Town, South
Africa.
Can. J. Zool. 77: 530–534 (1999)
berg and Rayner 1987; Fenton 1990; Barclay and Brigham
1991). Large species have high wing loading, which requires
that individuals fly rapidly to stay aloft, and means that they
are relatively unmaneuverable (Norberg and Rayner 1987).
They thus tend to forage in open areas. Such bats require
long-range prey detection and can achieve this by using
intense low frequency echolocation calls (Barclay and Brig-
ham 1991). Smaller species can fly more slowly and are
more maneuverable. They can thus forage in closer proxim-
ity to vegetation and can use higher frequency calls which,
although they do not provide as large a prey-detection range,
do provide greater information regarding the structural detail
and Stein 1980; Barclay and Brigham 1991).
While between-species comparisons support the body size –
habitat clutter hypothesis regarding the evolution of varia-
tion in echolocation call design (Barclay 1986; Aldridge and
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Barclay et al.
Rautenbach 1987; Fullard et al. 1991), within-species varia-
tion in echolocation call design also occurs. Individuals alter
the nature of their calls depending on the habitat (amount of
clutter) and the task at hand (Obrist 1995). Geographic vari-
ation between populations of the same species has also been
documented (Thomas et al. 1987), although it is not clear
whether such variation is due to body size or habitat differ-
ences between locations. A common problem in such com-
parisons has also been that recording and sound analysis
methods have differed between studies, potentially adding to
or obscurring natural variation in call features.
The purpose of our study was to test predictions based on
the body size – habitat clutter hypothesis using a geographi-
cally widespread species and a consistent recording and
analysis protocol. To do this we studied the echolocation
calls of the hoary bat (Lasiurus cinereus) because of its wide
geographic distribution and variation in body size across
subspecies.
Study species
The hoary bat is found throughout much of North and South
America, as well as on the Hawaiian and Galapagos islands
(Jacobs 1996; McCracken et al. 1997; Shump and Shump 1982).
Wherever it has been studied, L. cinereus is insectivorous and cap-
tures prey in the air. It uses a relatively low frequency, frequency-
modulated echolocation call (Belwood and Fullard 1984; Barclay
1986; Obrist 1995; McCracken et al. 1997). However, there has
been some debate (e.g., see McCracken et al. 1997) over the simi-
larity or difference between the calls used by Lasiurus cinereus
semotus in Hawai′i and those used by Lasiurus cinereus cinereus in
North America. Belwood and Fullard (1984) considered that the
two subspecies used similar calls, based on values for North Amer-
ica reported by Fenton and Bell (1981). Later, more detailed re-
cordings, however, suggested that L. c. cinereus used calls that differ
significantly from those of L. c. semotus (Barclay 1986). Again
differences in recording and analysis techniques or habitat may
have been involved. We thus analyzed calls recorded in the same
way from both subspecies.
Lasiurus cinereus semotus in Hawai′i is considerably smaller
(female body mass, 18.0 g) and has a lower wing loading (9.9 N·m–2)
than does L. c. cinereus from North America (34.0 g, 14.7 N·m–2;
Jacobs 1996). We thus predicted that mainland bats would use
lower frequency calls than the Hawaiian individuals.
Study sites
We recorded free-flying foraging L. cinereus at one site in Man-
itoba, Canada, in 1983 and 1984 (see Barclay 1986 for details) and
four sites in the Hawaiian islands: Koke′e State Park, Kaua′i
(1987); Kalopa State Park, Hawai′i (1987); Bradshaw Airforce
Base (= Pohakuloa; Jacobs 1996), Hawai′i (1992); and Ocean
View Estates, Hawai′i (1992). These sites varied in terms of the
density of the vegetation and the proximity that the bats flew to
this clutter. We could thus test predictions based on hypothesized
variation in echolocation calls due to habitat type and geographical
location.
In Manitoba, bats were recorded as they flew 5–15 m above the
ground (below tree canopy height) along a 5–10 m wide roadway
bordered by deciduous trees (Barclay 1986). Recordings at both
Koke′e and Bradshaw were of bats flying in the open around large
security lights. The bats flew 15–30 m above the ground and away
from (>20 m) vegetation (see also Belwood and Fullard 1984;
Jacobs 1996). We recorded bats at Kalopa and Ocean View as they
foraged in more cluttered situations. They flew 3–15 m above the
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531
ground, well below the canopy, and often within a few metres
of the trees surrounding the foraging areas. At Ocean View, we
recorded bats in two situations: as they flew over a 5–10 m
wide roadway and as they flew within the vegetation (′ohi′a
(Metrosideros polymorpha) forest) bordering the road (see also
Jacobs 1996). We analyzed the recordings in the two Ocean View
sites separately so that we could examine variation in echolocation
calls within the same population, but in two habitats differing in
amount of clutter. We predicted that bats foraging within the vege-
tation would use shorter higher frequency calls.
Based on the relationship between foraging habitat and echolo-
cation call design, we predicted that bats recorded at the more open
sites in Hawai′i (Koke′e and Bradshaw) would use lower frequency
calls than would those recorded in the more cluttered sites (Kalopa
and Ocean View). Geographic variation between populations on
different islands would be evident in the calls made by bats at the
two open sites (Bradshaw, Hawai′i, and Koke′e, Kaua′i).
Materials and methods
We recorded echolocation calls from all bats onto a Racal Store
4DS instrumentation tape recorder operated at 76 cm·s–1 using one
of three microphone systems: a Bruel and Kjaer (B and K) type
4135 (¼ in.; 6.3 mm) condenser microphone and type 2204 mea-
suring amplifier (Koke′e, Kalopa), a Larson–Davis type 2520
(¼ in; 6.3 mm) condenser microphone and type 2200C preampli-
fier (Ocean View, Bradshaw), and a custom-made condenser micro-
phone and power supply (Ocean View, Manitoba). The bandwidth
for the microphones and tape recorder in all situations was 5–
80 kHz (±2 dB). To verify the sensitivity characteristics of the cus-
tom-made microphone in the Ocean View site, we simultaneously
recorded bats with this microphone and the Larson–Davis micro-
phone. Frequency analyses of the calls revealed no spectral devia-
tions (±2 dB within the flat range) between the two systems.
All echolocation calls were analyzed in the laboratories of J.H.F.
using customized fast Fourier transform (FFT) spectral analyses
software, that employs a Cooley–Tukey algorithm and has a
minimum time and frequency resolution of 10 µs and 800 Hz (J.
Dawson, Queens University; personal communication) calibrated
with a Nicholet 100A minianalyzer.
As bats change the frequency and duration of their calls when
they search for, detect, and approach targets (Kick and Simmons
1984), we chose an arbitrary method to standardize which of the
calls that we analyzed. We only analyzed search phase calls, which
we defined as calls with at least 100 ms separating them from an-
other call. As these were recorded from wild bats, we could not
identify individuals and we have thus likely analyzed more than
one call per bat. We did, however, choose only one call from each
bat pass by the microphone. At all sites, at least six individuals
were present at any one time on a given night and we recorded
calls on more than one night at each site. Nonetheless, variation of
calls within a site was undoubtedly less than if we had been able to
analyze only one call per bat.
We measured a number of echolocation call variables. We used
the frequency with the highest intensity as a reference and deter-
mined higher and lower frequencies of calls at specific intensities
below the peak (Fullard et al. 1993). This eliminates the subjectiv-
ity involved in attempts to determine the minimum and maximum
frequencies in a call, which are often difficult to measure owing to
background noise. For each call, we measured duration in millisec-
onds, peak frequency in kilohertz (spectral peak of highest inten-
sity), Low18 (lowest frequency, 18 dB below peak), Low6 (lowest
frequency, 6 dB below peak), High18 (highest frequency, 18 dB
above peak), and High6 (highest frequency, 6 dB above peak). All
values were computed by one person (J.H.F.), who was unaware of
the identity of any of the recordings. Statistical analysis was also
conducted without knowledge of the locality. Call variables were
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532 Can. J. Zool. Vol. 77, 1999

Table 1. Characteristics of the echolocation calls of Lasiurus cinereus recorded from various locations.

Site Openness Duration (ms) Low18 (kHz) Low6 (kHz) Peak (kHz) High6 (kHz) High18 (kHz)
Bradshaw, Hawai′i Open 7.38 (1.78)a 26.15 (1.78)a 27.78 (1.91)a 29.81 (2.66)a 34.74 (4.84)a 45.46 (6.17)a
Koke′e, Kaua′i Open 7.69 (1.98)a 25.51 (1.29)b 27.02 (1.41)b 27.88 (1.70)b 32.04 (2.52)b 41.69 (4.72)b
Ocean View, Hawai′i Closed 9.63 (1.28)b 23.86 (1.27)c 25.26 (1.21)c 26.21 (1.16)c 29.57 (2.19)c 37.77 (3.46)c
Kalopa, Hawai′i Closed 10.27 (1.35)b 24.17 (1.18)c 25.79 (1.17)c 26.80 (1.14)c 32.28 (2.44)b 43.56 (3.65)a,b
Manitoba, Canada Intermediate 9.77 (2.52)b 18.30 (1.93)d 19.35 (2.06)d 20.11 (2.19)d 21.93 (3.28)d 27.73 (9.34)d
Note: Sample size was 100 calls for each site. Openness refers to our subjective ranking of how close the bats were flying to the vegetation and other
objects. Values are given as the mean with SD in parentheses. Within columns, means followed by different letters are significantly different (ANOVA with
Tukey’s test, p < 0.05).

Table 2. Comparison of echolocation call characteristics of Lasiurus cinereus flying in the open versus in the vegetation at Ocean
View, Hawai′i.
Duration (ms) Low18 (kHz) Low6 (kHz) Peak (kHz) High6 (kHz) High18 (kHz)
Open 8.92 (1.45)*** 24.72 (1.79)** 26.19 (1.76)*** 27.43 (2.24)*** 30.98 (3.18)ns 39.61 (5.08)**
Vegetation 5.46 (2.31)*** 25.90 (3.05)** 27.93 (2.22)*** 29.24 (2.80)*** 32.60 (4.35)ns 42.36 (6.31)**
Note: Sample size was 55 calls in the open and 69 calls in the vegetation. Values are given as the mean with SD in parentheses. Samples were tested
for differences using either two-tailed t tests or Mann–Whitney tests. ***, p < 0.001; **, p < 0.01; ns, p > 0.05.

analyzed for differences among locations using ANOVA and
Tukey’s tests. Calls made at Ocean View in the vegetation and in
the open were compared using t tests, or Mann–Whitney tests
when variances were unequal (F ratio test). All statistical tests
used α = 0.05.
being longer, calls made in the open had a narrower average
bandwidth (14.9 kHz) compared to those used in the vegeta-
tion (16.46 kHz).

Results
As we predicted, there were significant differences in all
call characteristics for bats recorded in different locations
(F[4,495] > 51.0, p < 0.0001 in each case; Table 1). The calls
of bats from Manitoba were significantly different in all fre-
quency variables from all other calls. They were of a lower
frequency in each case, peak frequency (the most reliable
measure) being 20.1 compared with 26.2–29.8 kHz for the
various Hawaiian populations. The Manitoba calls also had a
shallower frequency sweep (average bandwidth 9.4 vs. 13.9–
19.4 kHz). Call duration was no different from that of the
closed habitat populations on the Hawaiian islands. For repre-
sentative sonographs of calls from Manitoba and Hawai′i,
see Barclay (1986) and Belwood and Fullard (1984), respec-
tively.
Within Hawai′i, bats foraging in the two closed sites on
Hawai′i (Ocean View and Kalopa) used similar calls (Ta-
ble 1). Duration and three of the five frequency variables
were not significantly different. Calls at Kalopa started at
higher frequencies, although this is often the least reliable
measurement as atmospheric attenuation is greater at higher
frequencies (Griffin 1971).
Calls from bats recorded at the two open sites (Bradshaw
and Koke′e) on the different islands were significantly dif-
ferent in all frequency components (Bradshaw calls were
higher), but were not different in duration (Table 1). Calls at
these two sites were, however, more similar to one another
than they were to those recorded at the two closed sites. Bats
at the open sites had significantly shorter higher frequency
calls.
As predicted, bats flying within the vegetation at Ocean
View used significantly shorter higher frequency calls than
did those foraging in more open spaces (Table 2). Despite
Discussion
The echolocation calls of the free-flying hoary bats that
we recorded varied from location to location in most, but not
all cases, in accordance with our predictions based on varia-
tion in body size and habitat structure. Our prediction that
the larger subspecies from the mainland would use lower
frequency calls compared with those used by the Hawaiian
bats is supported by our data. Hoary bats from Manitoba
used significantly lower frequency calls compared with the
bats we recorded at all the Hawaiian sites. This confirms
earlier analyses (Barclay 1986; Obrist 1995) and qualitative
assessments (Fenton et al. 1983) from other North American
sites, and corrects the misconception that the two subspecies
use similar calls (Belwood and Fullard 1984; McCracken et
al. 1997).
North American hoary bats are almost twice as heavy as
their Hawaiian counterparts and have a significantly higher
wing loading (Jacobs 1996). This should mean that North
American individuals must fly faster to stay aloft and are
less maneuverable than Hawaiian individuals (Norberg and
Rayner 1987). Hawaiian hoary bats do fly in more cluttered
habitats than do North American individuals (Barclay 1985;
Jacobs 1996). The greater prey-detection range afforded by
lower frequency calls (Barclay and Brigham 1991) should
be less important to Hawaiian bats, which can react more
quickly to prey detected at short range. The improved detec-
tion of small prey, afforded by higher frequency calls, may
have been an important selection factor in the evolution of
the echolocation system of the smaller Hawaiian subspecies.
Lasiurus cinereus semotus is believed to have originated
from North American stock and has been present on the is-
lands for up to several million years (Morales and Bickham
1995). It is interesting that another island population of
hoary bats, on the Galapagos islands, is also composed of

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Barclay et al.
small individuals (17 g) that use calls similar in design to
those used in Hawai′i (29–22 kHz; McCracken et al. 1997).
On the Hawaiian islands, hoary bats foraging in the same
habitat used similar call designs. Bats foraging in the rela-
tively closed sites on Hawai′i (Ocean View and Kalopa) used
calls that were not significantly different in four of the six
variables we measured. Again the least reliable variables
(High6 and High18) were different. Likewise, calls from the
two open sites were more similar to one another than they
were to calls from either of the closed sites.
Between islands, there was some evidence for slight, but
statistically significant differences in call design. For exam-
ple, there were statistically significant frequency differences
between the calls recorded at the open sites on Kaua′i and
Hawai′i. The sites were similar in that the bats at both for-
aged high above the ground near bright lights and well away
from vegetation or other obstacles. Thus habitat differences
are unlikely to have influenced call design. Geographic dif-
ferences in echolocation call structure of bats have been
documented before (Thomas et al. 1987). It is not known to
what degree the populations of L. c. semotus on the various
Hawaiian islands are isolated, but our results suggest that
there may be slight echolocation call divergence.
Our analyses of the calls from bats foraging in different
habitats are somewhat contradictory. As predicted based on
the relationship between habitat clutter and echolocation call
design (Aldridge and Rautenbach 1987; Fenton 1990), bats
at Ocean View used shorter higher frequency calls when for-
aging in the vegetation than when foraging in the open.
Shorter calls may be necessary to avoid pulse–echo overlap
(Kalko and Schnitzler 1989) in situations where the bats fly
close to obstacles, and higher frequency calls, giving more
detailed information regarding small targets, can be used
where long-range prey detection is not a priority.
While the above data from a single site fit the predictions,
bats foraging at different sites behaved exactly opposite to
the predictions. Bats foraging at the two open sites used
shorter higher frequency calls than did those foraging at the
two closed sites. This is despite the fact that hoary bats at
the open sites in Hawai′i fly faster and are less maneuverable
than bats at the closed sites (Jacobs 1996), and should thus
have benefited from longer range calls. Although the echolo-
cation call differences were relatively small (approximately
2 ms and 2 kHz), they were consistent.
Explaining this apparent contradiction is difficult. Brad-
shaw and Ocean View provide the best comparison as the
sites are on the same island and were studied at the same
time and with the same equipment, and data on bat behav-
iour and insect abundance exist for both (Jacobs 1993). It is
unlikely that bats at Bradshaw and Ocean View were drawn
from different populations with different echolocation call
designs, given the relatively small size of the Big Island and
the ability of bats to move over large distances. Indeed, there
is genetic evidence that bats move between the study sites
(D.S. Jacobs, unpublished data). Differences in recording
conditions are also an unlikely explanation. Although bats at
Bradshaw were often further from the microphone than bats
at Ocean View, this would only serve to bias the recordings
at Bradshaw in favour of low frequencies, owing to the at-
mospheric attenuation of high frequencies (Griffin 1971).
Thus if anything, the differences we observed were conser-
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533
vative. Finally, it is possible that the light poles at Bradshaw
acted as clutter (see Kalko and Schnitzler 1993). This seems
unlikely to have affected the calls to the degree that we
found, however, as the bats at Bradshaw still foraged well
away from the poles (>10 m) most of the time.
Bats flying parallel to linear habitat edges, such as those
at Ocean View, may perceive the habitat as being unclut-
tered in front of them (E. Kalko, personal communication).
In this case, individuals may use narrower band calls than
expected. However, this would not explain the fact that calls
at Ocean View were more narrow and lower in frequency
than in those used at Bradshaw.
Different prey density at the open and closed sites is one
possible factor that could have influenced call design. If
concentrations of insects attracted to the lights at both open
sites meant that prey density was higher than at the closed
sites, foraging bats would encounter prey at a greater rate
and at a shorter range. In this case, shorter calls might be
used to eliminate pulse–echo overlap and higher frequencies
could be used, as maximizing prey-detection range would be
less critical. Prey density, however, is actually lower at
Bradshaw than at Ocean View (Jacobs 1993). On this basis
alone, encounter rates at Bradshaw should have been lower
and longer range calls more beneficial.
Prey size, bat flight speed, and mode of prey detection
may help explain the results. The insect fauna at Bradshaw
is dominated by large moths and the bats there prey heavily
on them. At Ocean View, a variety of small insects make up
the fauna and the diet of bats (Jacobs 1993). Despite the low
prey density at Bradshaw, the large target size and greater
echo strength combined with the greater flight speed of the
bats may mean that the detection rate was actually higher
at Bradshaw than at Ocean View. Indeed, our qualitative
assessment of the recordings indicated that Bradshaw had a
higher rate of terminal feeding buzzes (i.e., the rapid series
of calls produced as a bat attacks an insect; Griffin 1958)
than did Ocean View, despite a lower insect density. In addi-
tion, we suggest that hoary bats at Bradshaw (and Koke′e)
may detect and initially track large moths visually in the
bright lights. If so, echolocation may only be used to track
prey during the latter stages of the approach, thereby ex-
plaining the shorter duration and higher frequency calls.
Video analysis with simultaneous echolocation call record-
ings might help elucidate the interaction of visual and acous-
tic prey detection, and explain echolocation call variation
that is otherwise perplexing. In the end, however, corre-
lational studies such as ours will need to be supported by ex-
perimental studies testing the various explanations.
Acknowledgements
We thank the many people who assisted with the field and
laboratory work including L. Freed, C. Stone, D. Stone, Q.
Tomich, and C. Belanger, as well as the 150th Air National
Guard (Koke′e State Park) and those at the University of
Manitoba Field Station. Comments and suggestions by E.
Kalko and an anonymous reviewer significantly improved
the manuscript. Funding for our studies was provided by the
Audubon Society (Hawai′i chapter), the Hawaiian Division
of Forestry and Wildlife, U.S. Fish and Wildlife, the Na-
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534
tional Geographic Society, Sigma Xi, and the Natural Sci-
ences and Engineering Research Council of Canada.
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