Long- versus short-range foraging strategies of hoary (Lasiurus cinereus) and

silver-haired (Lasionycteris noctivagans) bats and the consequences for prey selection
ROBERTM. R. BARCLAY'
Zoology Department, University of Manitoba, Winnipeg, Man., Canada R3T 2N2
Received April 1 1 , 1985

BARCLAY, R. M. R. 1985. Long- versus short-range foraging strategies of hoary (Lnsiurus cinereus) and silver-haired
(Lasionycteris noctivagans) bats and the consequences for prey selection. Can. J . Zool. 63: 2507 - 25 1 5.
Habitat use, temporal activity, foraging behaviour, and prey selection of hoary bats (Lasiurus cinereus) and silver-haired
bats (Lasionycteris noctivagans) were studied at Delta Marsh, Manitoba. Bat activity was assessed by monitoring echolocation
calls with ultrasonic detectors. Prey availability was determined using sticky and Malaise traps and dietary information was
obtained from fecal analysis. Both species were active all night and foraged primarily in the lee of a narrow forested ridge.
Lasionycteris noctivagans foraged in a manner that indicates that it detects and pursues prey over short distances. These bats
Can. J. Zool. Downloaded from www.nrcresearchpress.com by UNIV CHICAGO on 02/22/13

fly slowly, are highly manoeuverable, and were commonly observed feeding on swarms of insects in small clearings. They
use echolocation calls that support the notion of a short-range foraging strategy and feed opportunistically on whatever insects
are available. Lasiurus cinereus, on the other hand, uses a long-range prey detection and pursuit foraging strategy. They fly
rapidly along straight line paths in open areas and use echolocation calls designed to detect insects at a distance. The diet
consists primarily of large insects (moths, beetles, and dragonflies), but the bats nonetheless feed opportunistically. The
foraging strategy likely restricts the availability and profitability of small insects as prey.

BARCLAY, R. M. R. 1985. Long- versus short-range foraging strategies of hoary (Lasiurus cinereus) and silver-haired
(Lasionycteris noctivagans) bats and the consequences for prey selection. Can. J . Zool. 63: 2507 - 25 15.
L'utilisation de l'habitat, I'activitk en fonction du temps, le comportement de recherche de nourriture et la sklection des
proies ont fait l'objet d'une ktude chez la chauve-souris cendrke (Lasiurus cinereus) et chez la chauve-souris argentke
(Lasionycteris noctivagans) a Delta Marsh, Manitoba. Des dktecteurs d'ultra-sons ont servi a enregistrer les cris d'kcho-
location. La disponibilitk des proies a ktk dkterminke au moyen de pieges collants et de pieges Malaise et I'analyse des feces
a permis d'ktudier le rkgime alimentaire des chauves-souris. Les deux eseces se sont avkrkes actives toute la nuit et elles
recherchaient leur nourriture dans la zone sous le vent d'une cr6te ktroite et boiske. Lasionycteris noctivagans semble dktecter
For personal use only.

la presence de ses proies et chasser sur de courtes distances. Ces chauves-souris volent lentement, font preuve d'une grande
manoeuvrabilitk et se nourrissent souvent au sein de nukes d'insectes dans les petites clairieres. Leurs cris d'kcholocation
indiquent qu'elles utilisent une stratkgie de recherche de nourriture de courte portke et qu'elles ont un comportement alimentaire
opportuniste, se nourrissant de tout insecte disponible. En revanche, Lasiurus cinereus fait de la dktection de longue portke
et poursuit ses proies sur de longues distances. Les chauves-souris de cette espece ont un vol rapide en ligne droite dans les
rkgions dkgagkes et leurs cris d'kcholocation leur permettent de dktecter leurs proies sur de longues distances. Leur rkgime
alimentaire est surtout constituk d'insectes de grande taille (papillons de nuit, colkopteres et libellules), mais leur mode
d'alirnentation derneure tout de m6me opportuniste. La mkthode de recherche de nourriture de ces chauves-souris rend
probablernent moins profitable la consommation des petits insectes, tout en rkduisant leur disponibilitk.
[Traduit par le journal]

Introduction
The foraging strategies employed by organisms are of inter-
est from evolutionary and ecological points of view. Differ-
ences in species' foraging strategies, evolved through inter-
specific competition and selection for resource partitioning,
may be important in determining community structure (Strong
et al. 1984). In an ecological sense, optimal foraging theory
predicts that natural selection should favour individuals capable
of choosing prey so as to optimize some aspect of energy or
nutrient intake or time output (e.g., Krebs 1978). In terms of
diet, morphological or physiological adaptations may restrict
the prey available to a species (e.g., Freeman 1979; Schmitt
and Holbrook 1984) and passive prey selection may result
since an individual is restricted in what it can consume. On the
other hand, prey selected by individuals for optimal foraging
reasons (e.g., Elner and Hughes 1978; Zach 1979) involves
active selection. Foraging strategies presumably involve a
combination of these two forms of prey selection.
Various aspects of foraging strategies have been studied in
insectivorous bats. Certain species are adapted for a gleaning
mode of prey capture (e.g., Fenton and Bell 1979; Bell and
'Present address: Biology Department, University of Calgary,
Calgary, Alta., Canada T2N 1N4.
Fenton 1984) or an ambush foraging strategy (e.g., Vaughan
1977), but most are aerial insectivores, catching prey on the
wing. Among aerial insectivorous species, one of the main
differences in foraging strategy may be between species that
detect insects at short range and those that detect them at long
range (Fenton and Bell 1979). This may have important im-
plications for other aspects of foraging strategies, including
prey selection.
There is much debate as to whether insectivorous bats select
certain types of prey or are opportunistic feeders. Many studies
indicate that insectivorous bats specialize on certain taxa (e.g.,
Ross 1967; Black 1972, 1974, 1979; Belwood and Fenton
1976; Buchler 1976; Goldman and Henson 1977; Belwood and
Fullard 1984; Warner 1985), body sizes (Ross 1967; Funakoshi
and Uchida 1975; Black 1979; Belwood and Fullard 1984), or
hardness (Freeman 1979, 198 1 ) of insects. Other studies,
however, indicate that opportunistic feeding is important
(e.g., Bradbury and Vehrencamp 1976; Fenton and Morris
1976; Gould 1978; Bell 1980) if only at certain times (Anthony
and Kunz 1977; Swift et al. 1985).
In studies reporting prey selection, prey availability has
often not been assessed (e.g., Ross 1967; Black 1972; Warner
1985) or is reflected in the diet (Black 1974). In other cases,
prey numbers rather than biomass have been analyzed (Buchler
 2508 CAN. J. ZOOL. VOL. 63, 1985
1976; Belwood and Fullard 1984). Determining prey avail-
ability is not easy, especially given the biases of insect sam-
pling techniques. In addition, various morphological, phys-
iological, and echolocation call characteristics of a bat species
may restrict what prey is actually available to it.
The purpose of this study was to investigate the foraging
strategies and prey selection of two sympatric aerial in-
sectivorous bats, the hoary bat (Lasiurus cinereus) and the
silver-haired bat (Lasionycteris noctivagans). In particular, I
was interested in the consequences of short- versus long-range
strategies on other aspects of foraging by these bats. Evidence
and preliminary observations suggested that L. cinereus is a
long-range insect detector and pursuer. It has one of the highest
wing loadings and aspect ratios of any species in North
Can. J. Zool. Downloaded from www.nrcresearchpress.com by UNIV CHICAGO on 02/22/13
America (Farney and Fleharty 1969) and is thus adapted for fast
but unmanoeuverable flight, a suggestion supported by obser-
vations (Hayward and Davis 1964; Belwood and Fullard 1984).
Lasionycteris noctivagans, on the other hand, has a lower wing
loading and aspect ratio (Farney and Fleharty 1969) and flies
more slowly but with more manoeuverability (Whitaker et al.
1977). These facts suggest that L. noctivagans is adapted for
short-range insect detection and pursuit.
Both L. noctivagans and L. cinereus are migratory, solitary,
tree-roosting species about which relatively little is known
(Barbour and Davis 1969; Kunz 1982; Shump and Shump
1982). Both species are widely distributed throughout North
America. The hoary bat is also found in Central and South
America and the Hawaiian islands. Lasiurus cinereus is one of
For personal use only.
the largest species in North America (adult mass 20-35 g;
Barbour and Davis 1969; Shump and Shump 1982) and re-
portedly preys heavily on moths (Ross 1967; Black 1972, 1974;
Belwood and Fullard 1984). Black (1974) considered L. cin-
ereus a "moth strategist" (i.e., specialist). Lasionycteris noc-
tivagans is a smaller species (8- 1 1 g) (Kunz 1982), and
although Black (1974) classed it as a moth strategist, a number
of other studies indicate it is more opportunistic (Whitaker
1972; Whitaker et al. 1981); a clear picture of the diet of this
species is not yet available.
The study area
The study was conducted in the vicinity of the University of Man-
itoba Field Station, Delta Marsh, Manitoba, Canada, located at the
southern end of Lake Manitoba (50'1 1 ' N; 98'23' W). Lake Manitoba
is a large, shallow lake that is separated from Delta Marsh by a narrow
(average width 80 m) forested dune ridge which runs for over 20 km
around the southern end of the lake. The ridge is dominated by
willows (Salix interior and S. amygdaloides), green ash (Fraxinus
pennsylvanica), and Manitoba maple (Acer negundo) (see Mackenzie
1982 for details). The marsh is composed of open bays, narrow
channels, and meadows. Emergent vegetation is dominated by the
giant reed (Phragmites australis) with smaller amounts of bulrush
(Scirpus spp.) and cattail (Typha spp.), while the meadows are dom-
inated by sprangle-top (white-top) (Scolochloafestucacea) and thistle
(Sonchus spp.).
Ice on the lake and marsh typically breaks up in late April or early
May and reforms in November. Prevailing winds, which can be very
strong because of the openness of the area, are from the northwest and
southeast and the forested ridge thus provides a windbreak.
Both the marsh and lake are extremely productive and insect abun-
dance is very high. 'Throughout the summer synchronous hatches of
chironomids occur and these dominate the insect fauna and form the
basis of the diet of the insectivorous birds nesting in the area (e.g.,
Busby and Sealy 1979; Sealy 1980).
Four species of bats are known from the Delta Marsh area, the little
brown bat (Myotis lucifugus), the red bat (Lasiurus borealis), the
hoary bat (L. cinereus), and the silver-haired bat (Lasionycteris noc-
tivagans) (Barclay 1 984). Only hoary and silver-haired bats are
common and present throughout the summer (Barclay 1984).
Methods and materials
The study was conducted during August and September 1981 and
May through September 1982- 1984. Roosting bats were located vi-
sually during the day by searching suitable sites in the forested ridge.
Lasiurus cinereus usually roosts high in tree foliage whereas L. noc-
tivagans roosts in small crevices in tree trunks (R. M. R. Barclay,
unpublished data). Bats were captured by hand from their roosts or in
Tuttle traps (Tuttle 1974) and mist nets as they foraged at night. Bats
were held overnight in individual cloth bags, sexed, aged as adult or
juvenile (young of the year) on the basis of ossification of the meta-
carpel phalangeal joints (Racey 1974), and banded on the forearm with
coloured plastic split rings (Barclay 1985). The bats were released on
the evening following capture and in 1981 their echolocation calls
were recorded using an Ampex tape recorder operated at 76 cm/s.
These calls were analyzed on a Kay 7029 sonograph.
Foraging activity was assessed in a number of ways. I made visual
observations of the departure of bats from roosts and of their foraging
behaviour. Foraging activity was monitored with ultrasonic micro-
phones to detect the echolocation calls of flying bats. 'This technique
has been used in a number of studies (e.g., Bell 1980; Fenton and
Thomas 1980) and allows identification of flying bats on the basis of
species-distinct echolocation calls. I monitored activity with a QMC
mini bat detector tuned to 35 kHz and a broadband ultrasonic micro-
phone coupled with an NLS MS-15 oscilloscope through a period
(zero crossing) meter (Simmons et al. 1979). The analysis of calls
recorded in 1981 from known individuals allowed me to identify to
species 97.6% (n = 5006) of the bat passes detected in the three
subsequent years.
Habitat and temporal aspects of bat activity were assessed along
three transects using the ultrasonic equipment. Transects were walked
or paddled at least once a week and included sections in the four main
habitats of the study area: along the beach on the north side of the ridge
(beach), along the south side of the ridge (ridge), through the marsh
meadows (meadow), and on the bays and channels of the marsh
(marsh). Each transect took approximately 1.5 h to traverse. Each
section of a transect was timed and the number of bat passes and
feeding buzzes was recorded. A feeding buzz is the rapid series of
echolocation calls produced by a bat as it attempts to capture an insect
and is easily identifiable on the bat detector.
During 1983 and 1984, temporal aspects of bat activity were inves-
tigated. A particular transect was traversed three times in a night, once
beginning 4 to 1 h after sunset, once ending 4 to I h before sunrise, and
once midway between the first and last.
At the beginning and end of each transect, ambient temperature
(+0.5'C), moonlight, cloud cover, precipitation, and wind speed
and direction were noted. Records of minimum and maximum tem-
peratures, continuously recorded wind speed and direction, and
precipitation were obtained from Environment Canada metereological
instruments present at the Field Station.
Aerial insect abundance and distribution was assessed at night
during 1982 and 1983 using sticky traps made from 38 cm long,
11 cm diameter black plastic pipe coated with Tanglefoot. Traps were
set at heights of 2 and 8 m on the beach and in a clearing on the south
side of the ridge, at 2 m in a meadow, and at 0 m on the lake and
marsh. Waves prevented placing high traps over the lake or marsh as
supporting posts would not remain anchored in the sediment. In 1982
additional traps were placed in the middle of the ridge and at 2- and
8-m heights on the south side of the ridge. In 1983, additional traps
were placed at 8 m in the meadow and at 2 and 8 m near a yard light
on the south side of the ridge where bats commonly fed. Comparisons
of insect abundance between years are based only on trap sites used
in both years.
Traps were positioned at sunset and retrieved at sunrise once a week
during 1982. In 1983, traps were used twice a week and positioned
during the same three time periods used for the transects.
 BARCLAY

TABLE1. Bat habitat use under different wind conditions

Mean passes/h ( k SD)

Species Wind Ridge Beach n t P
L . noctivagans North 18.362 18.83 3.09k7.87 31 4.34 <0.001
South
r 16 km/h 13.84k23.33 8.18k9.93 56 2.24 <0.05
>16 km/h 13.13k15.82 31.22531.26 33 3.50 <0.002
L. cinereus North 7.61 k 12.30 0.60k 1.99 11 T=6* <0.02
South
5 1 6 km/h 5.33k7.98 10.63513.18 32 2.38 <0.05
>16 km/h 3.2755.36 22.58k22.81 13 T=5* <0.005
NOTE:Paired comparisons from 1982 to 1984 are used; where sample sizes were small the Wilcoxon test was used (*).
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Insects were removed from the sticky traps and washed in EtOH to
remove any Tanglefoot adhering to them. Insects were identified to
order (family for dipterans) and counted and their body lengths were
measured to the nearest millimetre. After drying for 24 h at 95"C,
insects in each taxonomic group (and body length category for L . noctivagans
chironomids) were weighed to the nearest 0.0001 g.
In addition to the sticky traps, 1 m x 1 m Malaise traps made from
black cotton cloth were used to verify the insect trapping data. Malaise
traps were placed at 2 m on the beach, along the south side of the ridge
and in the meadow and at 0 m on the marsh. Because moderate to high
winds interfered with the use and efficiency of these traps and high
traps were not feasible, sticky trap results are used for most analyses.
The diet of L. noctivagans and L. cinereus was assessed by ana-
lyzing feces collected from captured bats, from under bat roosts, and
For personal use only.

from a gravel road above which L. cinereus commonly fed. Although

the validity of fecal analysis for determining the diet of insectivorous
bats has been questioned, it has been shown to give an accurate picture
(Kunz and Whitaker 1983). Ephemeroptera may be difficult to detect
in feces (Rabinowitz and Tuttle 1982) and thus the low proportion in
my study area simplified diet analysis. Much of the error in fecal
analysis comes from assuming that each pellet contains the remains of
one large or several small insects (Rabinowitz and Tuttle 1982). Since
I did not use numbers of prey in fecal samples, this source of error is
eliminated.
Fecal samples were stored individually, dried for 24 h at 95°C and
weighed to the nearest 0.0001 g. Individual pellets or fractions of
larger pellets were analyzed for insect remains by softening them in
70% EtOH and teasing them apart under a dissecting microscope.
Insect parts were identified using sticky trap specimens for reference,
and the percent volume contributed by each taxon in each pellet or
fraction was estimated visually to the nearest 5%. Data from the
pellets from each bat was pooled.
Information concerning the diet of L. cinereus was also obtained by
collecting culled insect parts dropped onto the gravel road above
which the bats fed. These were counted, measured, identified, and
compared with live specimens collected at a nearby ultraviolet light.
Various statistical tests are used to analyze the data. Analysis of
variance (ANOVA) and Student-Newman-Keuls tests were made
using SPSS programs. Other tests are from Zar (1984) and employ a
0.05 rejection criterion.
Results
Emergence and foraging behaviour
Both L. cinereus and L. noctivagans departed from their
roosts to feed approximately f h after sunset ( L . cinereus: n =
* *
37, mean SD = 35.2 6.6 min; L. noctivagans: n = 54,
* *
mean SD = 35.5 7.7 min). Weather conditions appeared
to affect the emergence of L. noctivagans. On three nights with
overcast conditions, all four bats that we monitored left their
roosts between 16 and 32 min after sunset. Nine L. noctivagans
remained in their roosts all night on two nights with ambient
L . cinereus
* 50
.c,
8-
F =10.6
.= -2 - p~0.0001
z2
'; 4-
-
I
. 80 26 125 93
B Ma R Me
FIG. 1 . Habitat use (mean passes per hour) by L. noctivafans (A)
and L . cinereus (B) using data from 1982 to 1984. The four habitats
are the beach or north side of the ridge (B), the south side of the ridge
(R), the marsh (Ma), and meadows (Me). Sample sizes (number of
transects) for each habitat are given. In this and subsequent histo-
grams, 1 SE is indicated and categories not significantly different
(Student-Newman-Keuls test) are indicated by a horizontal bar.
temperatures below 10°C and two remained in their roosts on
a night with heavy rain.
Both species were most commonly observed flying relatively
high above the ground near tree top level (ca. 10 m), although
other aspects of their foraging behaviour were quite different.
On numerous occasions L. noctivagans fed in small clearings
in the ridge. Individuals appeared to be highly maneuverable
and darted in and out of insect swarms. In contrast, Lasiurus
cinereus was most commonly observed flying in a straight
line fashion in open situations. Lasiurus cinereus were never
observed feeding on prey swarms.
Feeding buzz rates were strikingly different for the two
species. Three individual L. noctivagans that were observed
feeding continuously for 2 to 5 min each produced between 4.3
and 12.9 buzzes per minute. An average of 0.32 feeding buzzes
 CAN. J . ZOOL. VOL. 63. 1085

TABLE2 . Average ( k SD) nightly percent biomass contributed by various insect taxa in sticky trap
samples ( 1982 and 1983)

May June July Aug. - Sept .

(6 nights) (1 2 nights) (I 1 nights) ( I 1 nights)

Chironomidae
Lepidoptera
Culicidae
Muscoid flies
Tipulidae
Coleoptera
Trichoptera
Ephemeroptera
Homoptera
Hemiptera
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Neuroptera
Totals
- --

N O T ~The
: total number of each taxon caught IS shown In parentheses Mlnor amounts from other taxa made up the remalnlng blomass

L . noctivagans was true for L . noctivagans in all 3 years and for L . cinereus
@ I in 1982 and 1984. A small sample size and high variability in
1983 likely produced the nonsignificant result in that year.
For both species, activity levels were lowest over the
meadow in all 3 years while the greatest activity was either
along the beach or ridge. This was supported by feeding buzz
rates which were highest for both species over the ridge.
Wind speed and direction affected habitat use by both
For personal use only.

species (Table 1). Paired comparisons on the same night

L . cinereus
@ I F=1.72
FIG.2. Bat activity (passes per hour) during the three nightly sam-
pling periods (early (E), middle (M), and late (L)) in 1983 and 1984.
(A) L. noctivagans. (B) L. cinereus. n is the number of nights.
per pass were heard for L . noctivagans in 1982 along the
transects. In contrast, the mean feeding buzz rate for individual
L . cinereus was 1.02 ? 0.76 per minute (range 0.18- 3.80;
n = 30) and an average of only 0.12 buzzes per pass were
recorded in 1982.
Individual L . cinereus occasionally established what ap-
peared to be feeding territories, particularly around lights on
buildings or in the lee of the ridge during strong winds. These
individuals foraged back and forth along 40- to 50-m routes and
reacted to other bats by pursuing them, emitting audible vocal-
izations and occasionally coming in contact with them. The
pursued bat almost always disappeared after a chase. Lasionyc-
teris noctivagans was never observed acting in this way and
several individuals often fed in close proximity.
Neither species was active uniformly in the four habitats in
the study area, as indicated by the transect data (Fig. I). This
showed that L . cinereus was most active on the lee side of the
ridge in north or south wind conditions. Lasionycteris noc-
tivagans acted similarly (Table I), although only in strong
south winds (>16 km/h) did they shift from the south to the
north side of the ridge.
Although foraging activity for both species was greatest
during the middle time period (ca. 0030-0200), the temporal
patterns were not significantly different from random (Fig. 2).
The temporal activity of L . noctivagans was affected by ambi-
ent temperature. When the temperature during the middle time
period was greater than 14"C, activity was higher than during
the early time period (Wilcoxon test, T = 143.5, p < 0.05,
n = 31) while on cooler nights this trend was reversed, al-
though the difference between early and middle activity was
not significant ( T = 98.5, p > 0.05, n = 23).
Insect abundance and distribution
Over 35 000 insects (dry weight 27.8 g) were captured on the
sticky traps in 1982 and 1983 and, as expected, chironomids
dominated in both numbers and biomass. Although both spe-
cies of bats often foraged at heights somewhat above the sticky
traps, chironomids were found at all heights and were un-
doubtedly the dominant insects where the bats foraged. There
were considerable differences between the years, however.
Synchronous hatches of chironomids (particularly large species
(body length > 10 mm)) were smaller in 1983 (Fig. 3A),
possibly due to cold, dry spring conditions. There was no
noticeable difference in the nonchironomid biomass pattern
(Fig. 3B).
Chironomids were particularly abundant in June (Table 2),
averaging 8 1% of the nightly biomass on sticky traps and 67%
in Malaise traps. Chironomids were relatively less abundant at
other times during the summer but were still the largest com-
ponent of the insect biomass in July and August (Table 2). The
calculations use the mean nightly biomass percentages rather
 BARCLAY
All Insects
A
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1.0- @
A-A
Without Chironomids ,,1 982
983
,
-
0
w
U)
(II
U)
For personal use only.
I 0.5-
2 p = 0.67
n -
'a
FIG.3. Comparison of seasonal insect biomass from sticky traps in
1982 and 1983. (A) All insects; (B) without chironomids. The large
peak in mid-August 1983 was due to the capture of two large odo-
nates.
than the percent of the total monthly biomass since this reduces
the influence of those nights with extremely high chironomid
biomass.
Moths constituted an average of 12.8% of the nightly bio-
mass and < 1 % of the total number of insects caught on sticky
traps and 14.4% of the biomass and 3.1% of the number of
insects in the Malaise trap samples.
In addition to considerable seasonal fluctuation in insect
abundance (Fig. 3), low ambient temperatures in May and June
reduced insect abundance. Insect biomass was significantly
correlated with ambient temperature during the three time
periods in 1983 (one-tailed Spearman rank correlation, early
r, = 0 . 6 3 , ~< 0.05; middle r, = 0 . 6 5 , ~< 0.05; later, =
0.82, p < 0.01; all night r, = 0.77, p < 0.025; n = 9 nights).
Insect biomass was highest in the early time period, although
this difference was not significant unless one night with ex-
tremely high small chironomid catches is eliminated from the
data (Fig. 4). This difference was due to differences in chi-
ronomid and other small insect activity since the temporal
activity for large (body length > 10 mm), nonchironomid in-
sects was random ( F = 0.89, p = 0.42, df = 50). Nonetheless,
FIG.4. Temporal insect biomass ( 5SE) from sticky traps in 1983.
n is the number of nights of sampling.
Middle n =19
0.01-
I I
F = 1.94
p=0.13
Late n=17
0.01- 1 I
I : l I : I
R 6 Me Ma
FIG. 5. Distribution of sticky trap insect biomass by habitat (abbre-
viations as in Fig. 1) and time period in 1983. n is the number of nights
of sampling.
chironomids maintained their dominance of the biomass, never
making up less than 23% of the total in any time period of
any month.
In both 1982 and 1983 the highest mean insect biomass per
night occurred over the marsh, using the low traps for com-
parisons. The difference was only significant in 1983, how-
ever, and was due to high insect abundance over the marsh in
the early period (Fig. 5). Presumably this was due to syn-
chronous dusk emergence of adult insects from larval stages
inhabiting the marsh (e.g., Chapman 1971). Biomass was
equally distributed over the four main habitats in the middle
and late periods (Fig. 5).
Nonchironomid, large insect biomass was randomly distrib-
uted over the four habitats on a mean nightly basis ( F = 0.92,
p = 0.4.4, df = 71), although the ridge and beach had the
highest amounts.
Wind direction did not affect insect distribution along the
ridge since ridge and beach sticky traps had statistically equal
biomass in paired comparisons (south wind: t = 1.02, p >
 25 12 CAN. J . ZOOL. VOL. 63. 1985

TABLE3. Fecal analysis of 64 samples from L. noctivagans (monthly TABLE5 . Seasonal differences in L. cinereus diet (mean percent bio-
mean percent composition k SD) mass k SD) as determined by fecal analysis using feces collected
under roosts
May June Aug. -Sept. Total
(n=33) (n=21) (n=10) (n=64) June (n= 12) July (n=48) August (n= 14)
Chironomidae Lepidoptera
Muscoid flies Odonata
Coleoptera Coleoptera
Tipulidae Muscoid flies
Lepidoptera Culicidae
Culicidae Tipulidae
Neuroptera Trichoptera
Homoptera Neuroptera
Hemiptera Homoptera
Trichoptera Chironomidae
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Other+ Other+ unknown

unknown
and beetles with chironomids constituting a very low per-
TABLE4 . The diet (as mean percent biomass k SD) of L. cinereus centage (Table 4). Individuals ate different insects at different
determined by analyzing feces collected from under roosts, trapped times of the night and of the summer. Feces from bats trapped
bats, and the ground under feeding bats while foraging in the middle of the night, and from the ground
under foraging bats, contained high levels of moths and beetles
Roost Trap Ground Total and virtually no odonates. On the other hand, feces from under
(n=77) (n= 15) (n=9) (n= 101) roosts, representing the prey eaten by the bats just prior to
Lepidoptera roosting at dawn, contained high levels of odonates in addition
Odonata to moths and beetles (Table 4). Comparison of feces collected
Coleoptera under the roosts (Table 5) indicated that the diet was more
Muscoid flies diverse in June with a low odonate component, compared with
For personal use only.

Culicidae that in July and August.

Tipulidae
Trichoptera
Neuroptera
Homoptera
Chironomidae
Other+
unknown
0.05, n = 66, paired t-test; north wind: T = 54, p > 0.05,
n = 15, Wilcoxon test).
Insect biomass tended to be higher on 8- versus 2-m traps.
This was true for 1982 (paired t-test, t = 2.16, p < 0.05, n =
29) but only in May and June in 1983 (Wilcoxon test, ridge T
=30.5,~ < 0.001, n = 23; beach, T = 84,p = 0.02, n = 26).
The height difference was not evident for the entire 1983 data
set ( t = 1.02, p > 0.05, n = 81).
Diet
Since the size of fecal samples varied, I used the percent
composition of each sample (by biomass) when analyzing the
diet and calculated sample means. The main conclusions
reached using this method are no different from those reached
if percent of the total fecal biomass were used.
Fecal samples from 64 L. noctivagans were analyzed (dry
mass 2.44 g). The bats had fed on a variety of insects, primarily
dipterans and other small insects (Table 3), with muscoid flies
accounting for almost half the diet in May, chironomids dom-
inating in June, and a variety of insects being eaten in August
and September.
The size of the insect parts in L. noctivagans feces indicated
that small insects constituted the bulk of the diet. Insect wings
3- 10 mm in length were common as were eyes from 2 to
10 mm body length chironomids.
I analyzed 101 fecal samples from L. cinereus (dry mass
9.63 g). The diet was composed mainly of moths, odonates,
Although some insect wings as small as 3 mm in length were
found in L. cinereus feces, evidence suggested that larger in-
sects were important in the diet. Beetle remains included wings
over 10 mm in length and legs from Dytiscidae with body
lengths of about 15 mm. In addition, moth wings collected
from the ground under feeding bats averaged 17.1 mm in length
(n = 50, SD = 0.23, range 12-23 mm) and indicated moths
with body lengths of 14-29 mm were being eaten.
Discussion
The foraging strategies of L. noctivagans and L. cinereus are
similar in some respects, but the distinction between short-
and long-range strategies is supported by this study and likely
has important implications for habitat and prey selection.
Both species are aerial insectivores, depart from their roosts
relatively early compared with individuals in more southern
locations (e.g., Kunz 1973; Shump and Shump 1982), and are
active all night and in the same habitats. Thus, major temporal
and spatial partitioning of the resources is not apparent. Envi-
ronmental conditions affect where and when both species feed.
They feed primarily along the lee side of the ridge, possibly
selecting areas where flight or insect capture is more efficient.
Lasionycteris may be less affected by south winds because
their manoeuverability allows them to feed in small, slightly
sheltered clearings on the south side of the ridge, whereas
L. cinereus is restricted to more open areas because of its fast,
unmanoeuverable flight.
Foraging by L. noctivagans is reduced at low ambient tem-
peratures, possibly because low temperatures in the spring
reduce insect abundance. Similar observations have been
made for little brown bats (Myotis lucifugus) (Anthony et al.
1981) and suggest that prey becomes a limiting factor at low
temperatures.
Despite the above similarities, the two species have distinct
 BARCLAY
foraging strategies. Lasionycteris is a highly manoeuverable
bat that can efficiently prey on swarms of insects such as
chironomids and feeds on whatever insects are available. In this
study, the diet, composed primarily of small dipterans, does
not support the "moth strategist" label initially placed on
this species (Black 1974). In this and other studies, there is
seasonal, geographic, and individual variation in the diet
(Whitaker 1972; Whitaker et al. 1977; Whitaker et al. 1981),
supporting the notion of an opportunistic foraging strategy.
In this study, chironomids increased in importance in the
diet as they increased in relative abundance. In August and
September, however, chironomids made up a significant pro-
portion of the trap samples but were almost absent from the
diet. A similar seasonal shift in selectivity was noted for
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M. lucifugus (Anthony and Kunz 1977) and was explained as
active prey selection at times when prey were most abundant.
This cannot explain the shift here since what appears to be prey
selection occurred when insect abundance was low. Changes
in prey size may have been important. Whereas in June
large chironomids (>10 mm body length) made up 67.2%
(1982) and 26.8% (1983) of the insect biomass, in August and
September they accounted for only 12.8% (1982) and 7.3%
(1983). This may partly account for the reduced chironomid
component in the diet, since small insects are less detectable by
bats (see below).
Lasiurus cinereus is a fast, less maneuverable flier that
feeds primarily in open areas. Unlike L. noctivagans, indi-
vidual L. cinereus sometimes establish feeding territories
For personal use only.
(Belwood and Fullard 1984; this study), indicating that at times
prey is scarce enough and prey patches are important enough to
defend them. The flexibility in foraging behaviour indicates
that a degree of opportunism in the foraging strategy exists.
This study indicates that, contrary to several other studies
(Black 1972, 1974; Belwood and Fullard 1984), L. cinereus is
not a "moth strategist." Odonates and beetles are important in
the diet, supporting Freeman's ( 1981) contention that L. cin-
ereus has the jaw and tooth structure to eat hard-bodied insects.
Overall it appears that opportunistic feeding is important to
L. cinereus because there is dietary variation within a night
(this study), seasonally (this study), and geographically (Black
1972, 1974; Whitaker and Tomich 1983; Belwood and Fullard
1984).
Despite the opportunistic aspects of the L. cinereus foraging
strategy and diet, it is clear from this study that these bats do
not feed on insects in proportion to their abundance. What is
not clear is how important the long-range foraging strategy is
in producing prey selection by affecting prey availability and
profitability.
Individual Lasiurus cinereus could actively select prey for a
number of reasons. Some predators select prey so as to max-
imize net energy intake (e.g., Elner and Hughes 1978; Zach
1979) and this has been used to explain prey selection by some
insectivorous bats (Belwood and Fenton 1976). Such prey
selection is usually related to prey size and the explanation is
only satisfactory if pursuing and handling a small prey item is
more energetically costly than ignoring small prey in favour of
searching for and capturing larger prey. For insectivorous bats,
which detect insects via echolocation only at short distances
(Fenton 1982), search time is relatively long compared with
pursuit time and prey are handled rapidly (Kallen and Gans
1972). Thus, the cost of consuming small prey is relatively
small and I suggest energetically based prey selection for opti-
mal foraging reasons should only occur at extremely small prey
25 13
sizes, a suggestion also made by Swift et al. (1985). In other
words, bats should pursue all but the smallest insects they
detect.
Certain nutrients may be critically important for bats and
differences in the nutrient content of different prey types could
be the basis for active prey selection (e.g., Krebs 1978). Un-
fortunately there is very little information regarding the nutrient
content of different insects. Furthermore, for insectivorous bats
to actively select prey on this basis they must be able to dis-
tinguish among the various taxa. It is still not known whether
echolocation can provide enough information for bats to do this
(Fenton 1982).
For insectivorous bats, active prey selection, for whatever
reason, is easier to visualize on the basis of prey size and this
has been suggested in several studies (Ross 1967; Black 1979;
Belwood and Fullard 1984). Optimal foraging, however, is not
the only explanation for such selection.
Small insects such as chironomids are less available as prey
than their abundance indicates. Different sized insects are not
equally detectable to echolocating bats (see also Anthony and
Kunz 1977) since larger insects reflect more intense echoes, are
detectable at greater distances, and thus appear to a bat as more
abundant than they really are. This alone, however, can not
account for the lack of chironomids in the diet of L. cinereus
because in June a large percent of the insect biomass was due
to large chironomids (>10 mm body length).
Although body length has been used in the past as a measure
of prey size, to an echolocating bat the critical characteristic
related to prey detection is reflective surface area. Roeder
(1963) showed that the wings of moths were the important
feature influencing echo intensity and that the body played a
"negligible" role. Similarly, Kick (1982) showed that bats de-
tect large-diameter targets at greater ranges than small targets.
Since a moth has a wing area six times as large as a similarly
sized (body length) chironomid (R. M. R. Barclay, unpub-
lished data), moths and other fat-bodied, large-winged insects
will appear much more common than they really are while
chironomids and other thin-bodied, narrow-winged insects will
be detected less often than their abundance suggests.
Although the above arguments could be made for both
L. noctivagans and L. cinereus, adaptations for short- and
long-range foraging strategies may differentially influence prey
availability and profitability for the two species. The echo-
location calls used by L. cinereus to search for insects are
consistent with a long-range prey detection foraging strategy
and enhance the detectability of large insects (R. M. R.
Barclay, in preparation). The calls are low and essentially
constant in frequency, a structure adapted for detecting insects
in the open at long range (e.g., Simmons and Stein 1980;
Lawrence and Simmons 1982). The low frequency (20- 16
kHz), however, means that targets smaller than the wavelength
of the sounds (ca. 2 cm) reflect weaker echoes than larger
targets. Thus, very small insects (chironomids have wing
widths of 2 mm or less) may not be detected by L. cinereus or
are only detected at close range. The fast, unmanoeuverable
flight of L. cinereus means that insects detected only at close
range are less available as prey because there is insufficient
reaction time for the bats to manoeuver for the capture.
Swarming insects such as chironomids may also not be as
energetically profitable as for a more manoeuverable bat. Las-
iurus cinereus thus employs a style of flight and echolocation
calls that suggest it pursues individual prey detected at long
range, a foraging strategy which reduces the availability and
 25 14 CAN. J. ZOOL. VOL. 63, 1985
profitability of small and (or) swarming prey and may lead to
the diets observed in this and other studies.
Lasionycteris noctivagans, on the other hand, uses higher,
frequency-modulated echolocation calls (R. M. R. Barclay, in
preparation), which do not travel as far (Lawrence and
Simmons 1982) but are more efficient at detecting small
insects. When combined with the observations of slow,
manoeuverable flight, this species appears adapted to pursue
insects detected at short range and efficiently use swarming
insects as prey.
Although short- versus long-range foraging strategies appear
to be important features of the two species in this study, it
remains to be determined how much influence these strategies
have on habitat and prey selection by other aerial insectivorous
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bats.
Acknowledgements
I thank Melissa Todd, Dan Farr, Kevin Cash, Elizabeth
Straszynski, Gaynette Friesen, and Jean Dabros for their dedi-
cated assistance with field and laboratory work. M. Brock
Fenton kindly loaned me equipment and Dan Guinan, Gloria
Pohajdak, Heidi den Haan, James Briskie, and Spencer Sealy
located roosting bats during the day. The Portage Country Club
allowed me access to their property for this research and the
facilities of the University of Manitoba Field Station were made
available by Jennifer Shay. Tom Kunz, Brock Fenton, James
Fullard, Donald Thomas, Joseph Culp, Jackie Belwood, Gary
For personal use only.
Bell, and two anonymous reviewers provided enlightening
comments on the manuscript. The study was conducted while
the author was a Natural Sciences and Engineering Research
Council of Canada (NSERC) University Research Fellow and
was supported by NSERC operating and equipment grants as
well as a grant in aid of research from the University of
Manitoba. This is publication No. 135 of the University of
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