J.
Zool.,
Lond.
(1978)
186,
403-406
No es
on
he
d y
season
oos ing
and
o aging
beha iou
o
Epomopho us
gambianus
and
Rouse us
aegyp iacus
(Chi op e a
:P e opodidae)
DONALD
W.
THOMAS
AND
M.
BROCK
FENTON
Depa men
o
Biology,
Ca le on
Uni e si y,
O awa,
Canada
(Accep ed
11
Ma ch
1978)
(Wi h
1
igu e
in
he
ex )
We
s udied
he
mo emen s
and
o aging
beha iou
o
Epomopho usgambianus
and
Rouse us
aegyp iacus
du ing
he
d y
season
in
Rhodesia.
The
nigh ly
ac i i y
o
bo h
species
was
concen a ed
in
he
ipa ian
o es whe e
hey
ed
on
he
ui s
o
Diospy os
senensis.
Epomopho us
gambianus
oos ed
du ing
he
day
high
in
he
hick canopy
o
T ichelia
eme ica
o
Kigelia
a icana
along
he
i e
banks
and
began
o aging,
each
one
in
a
p e e ed
D.
senensis
sh ub,
be o e
1845
h s.
Rouse us
aegyp iacus
a i ed
in
he
s udy
a ea
app ox-
ima ely
one
hou
la e
and
appea ed
o
o age
in
g oups.
The
empo al
sepa a ion
o
he
eeding
imes
o
he
wo
species
o
ba s
may
p o ide
a
basis
o
he
pa i ioning
o
a
single
species
ood
esou ce.
In
spi e
o
nume ous
publica ions
on
A ican
ui
ba s
(e.g.
Bake
&
Ha is,
1957;
Rose ea ,
1965;
B osse ,
1966;
Jones,
1971,
1972;
Ayensu,
1974;
Kingdon,
1974),
he e
a e
ela i ely
ew
da a
conce ning
he
eeding
and
oos ing
beha iou
o
p e opodids
(see
also
B adbu y,
1977).
We
p esen
he e
some
obse a ions
on
o aging
beha iou
and
habi a
use
o
Rouse us
aegyp iacus
(Geo oy)
and
Epomopho us
gambianus
(Ogilby)
in
deciduous
woodland
du ing
he
d y
season.
Ou
obse a ions
we e
made
a
he
Hos es
Nicolle
Ins i u e
o
Wild
Li e
Resea ch
in
he
Sengwa
Wild
Li e
Resea ch
A ea,
ca.
110
km
wes
o
Gokwe,
Rhodesia
(18°10'S;
28°13'E)
be ween
7
and
28
June
1977.
We
sampled
se e al
habi a s
including
mopane
and
miombo
woodlands,
ipa ian
o es ,
and
a eas
o e
wo
la ge
i e s
(Sengwa
and
Lu ope),
using
9
o
12
m
long,
35
mm
mesh
mis
ne s.
Fo
de ails o
habi a
and
physiog aphy
see
Cumming
(1975).
A
o al
o
19
p e opodids
was
cap u ed,
including
10
E.
gambianus
(all
adul
emales)
and
nine
R.
aegyp iacus
(se en
adul
emales,
one
subadul
emale
and
an
adul
male).
Eigh een
o
he
ui
ba s
we e
ne ed
a
one
si e
along
he
Lu ope
Ri e
and
in
he
adjacen
ipa ian
o es ,
gene ally
in
he
icini y o
Diospy os
senensis
Ko zsch
(Ebenaceae)
sh ubs,
he
majo
ui
species
a ailable
du ing
ou
s udy
pe iod.
Diospy os
senensis,
a
all
sh ub
o
small
ee
es ic ed
o
he
a eas
along
i e s,
has
ui s
ha
ipen
synch onously
and
weigh
an
a e age
o
1
-9
g
when
husked.
Al hough
Balani es
maughamii
(Sp ague)
was
also
ui ing
du ing
ou
s udy,
i
was
nei he
widesp ead
no
common.
To
moni o he
mo emen s
o
indi idual
ba s
we
used
adio
ansmi e s
ha
we e
buil
in
he
Ca le on
Uni e si y
Science
Wo kshop
(8
g
po ed)
and
had
a
de ec ion
ange
o
2
km
in
la
e ain
and
11
km
om
a
high
an age
poin .
We
agged
wo
E.
gambianus
and
one
R.
aegyp iacus
by
gluing
he
ansmi e
o
he
mid-do sal
u .
Howe e ,
ou
da a
on
mo emen s
a e
o
E.
gambianus
alone
since
we
los
con ac
wi h
he
R.
aegyp iacus
403
404
D.
W.
THOMAS
AND
M.
B.
FENTON
a e
we
had
ollowed
i
5
km
om
he
elease
si e
on
he
i s
nigh .
Loca ional
ixes
we e
made
using
a
ou
elemen
Yagi
an enna
and
po able
ecei e .
Day ime
ixes
se ed
o
loca e
oos
si es,
and
noc u nal
ixes
(1845-1945
h s),
including
wo
hou s
o
con inuous
obse a ion
o
each
ba
(2000-2200
h s),
we e
used
o
assess
mo emen s
and
beha iou
associa ed
wi h
eeding.
Mo emen s
o
he
wo
agged
Epomopho us
we e
concen a ed
a ound
a
se ies
o
oos
and
Kigelia
ipa ian
sh ub
and
g asslandwi h
Diospy os
senensis
1-;-;';"-j;
Colophospe mum
mopane
woodland
(A);
Acacia
ipa ian
communi ies
(B);
and
B achys egia
boehmii-Comb e um-Hypa henia
wooded
and
bushed
g assland
(C).
NOTES
ON
FORAGING
OF
FRUIT
BATS
405
open
woodland
which
s e ched
abou
300
m
back
om
he
i e
banks
(Fig.
1).
On
he
six
days
ha
we
loca ed
oos s
o
E.
gambianus,
i e
di e en
ees
we e
used:
h ee
T ichilia
eme ica
(Chio .),
and
wo
Kigelia
a icana
(Lam.).
One
ba
used
he
same
T.
eme ica
on
wo
consecu i e
days
and
hen
mo ed
only
10
m
o
an
adjacen
T.
eme ica
on
he
hi d
day,
whe eas
he
o he
indi idual
mo ed
150
and
400
m
be ween
wo
K.
a icana
and
one
T.
eme ica
o e
he
h ee
days.
Bo h
animals oos ed
high
in
he
cen e
o
he
canopy,
10-15
m
abo e
he
g ound,
and
one
ba
ha
we
isually
loca ed
on
wo
days
oos ed
alone.
By
ixes
we
de e mined
ha
E.
gambianus
began
o aging
less
han
45 min
a e
da k
(dusk
a
1800
h s).
Each
ba
used
he
same
D.
senensis
sh ub
exclusi ely
on
bo h
nigh s.
Du ing
one
pe iod
o
con inuous
obse a ion,
one
ba
spen
105
min
in
i s
eeding
ee
and
15
min
in
an
adjacen
T.
eme ica
be o e
e u ning
o
he
D. senensis.
The
o he
ba
spen
90
min
in
i s
eeding
ee,
10
min
in
an
adjacen
K.
a icana,
and
20
min
again
in
he
D.
senensis.
Since
we
ne e
obse ed
o he
E.
gambianus
in
o
nea
hese
wo
eeding
ees,
we
sugges
ha
his
species
may
eed
alone.
Benea h
he
D.
senensis
sh ubs
we
ound
sca e ed
husks
o
ui s
bu
no
seeds,
indica ing
ha
he
ba s
had
p obably
consumed
he
en i e
ui s.
Coinciden
wi h
he
seden a y
eeding
beha iou
o
E.
gambianus,
we
obse ed
ew
lying
p e opodids
be ween
dusk
and
2000
h s.
A e
his
he e
was
a
ma ked
inc ease
in
he
ac i i y
o
p e opodids,
co esponding
o
he
a i al
o
Rouse us
which
we
iden i ied
by
hei
cha ac e is ic
clicking
sounds
p oduced
du ing
ligh .
We
knew
o
no
oos s
(ca es)
wi hin
a
leas
11
km
o
he
Lu ope
si e,
and
he
la e
a i al
o
he
Rouse us
was
appa en ly
due
o
he
ela i ely
long
commu ing
dis ances.
Rouse us
aegyp iacus
appea ed
o
o age
in
g oups
and
we
o en
obse ed
as
many
as
i e
indi iduals
lying
in o
and
om
a
single
D.
senensis
sh ub.
The
a i al
o
he
Rouse us
in
a eas
whe e
Epomopho us
we e
eeding
in
D.
senensis
sh ubs
did
no
al e
he
beha iou
o
he
la e
ba s,
and
we
common-
ly
wa ched
bo h
species
eeding
in
he
same
ui
sh ub.
The e
is
a
d ama ic
di e ence
in
species
composi ion
and
habi a
use
in
he
ui
ba
communi y
o
ou
s udy
a ea
be ween
we
and
d y
seasons.
The
smalle
Epomopho us
wahlbe gi
and
E.
c yp u us
also
occu
in
he
a ea
in
he
we
season,
bu
a
ha
ime
hese
species
and
he
E.
gambianus
a e
mo e
widesp ead,
occu ing
in
se e al
habi a s
(Fen on,
1975).
The
p esence
o
wo
simila -sized
ui
ba s
(100
o
130
g)
in
he
d y
season
using
he
same
single
species
ood
base
poses
an
in e es ing
ques ion
in
esou ce
pa i ioning.
Sympa ic
P e opodidae
o en
show
b oad
o e lap
in
habi a
use
and
ood
selec ion
(Bake
&
Ha is,
1957;
Rose ea , 1965;
Jones,
1971,
1972;
Smi he s,
1971;
Smi he s
&
Lobao
Tello,
1976).
Howe e
pa i ioning
o
ood
esou ces
by
hese
ba s
may
occu
h ough
ine
spa ial
o
lamina
sepa a ion
o
eeding
si es
wi hin
he
same
ees
(Jones,
1972).
Fo
example,
Eidolon
hel um
and
Epomops
anque i
a e
known
o
o age
in
he
same
ees
bu
a e
sepa a ed
spa ially
by
hei
ela i e
deg ees
o
pene a ion
in o
he
oliage;
he
la ge
E.
hel um
uses
he
pe iphe al
b anches
and
he
smalle
E.
anque i
mo es
deepe
in o
he
oliage
(Jones,
1972).
The
Epomopho us
gambianus
and
Rouse us
aegyp-
iacus
which
we
obse ed
in
he
same
sh ubs
a
he
same
ime
showed
no
such
spa ial
sepa a ion.
In
ou
s udy,
he
use
o
di e en
oos s
(ca es
s.
oliage)
a
di e en
dis ances
om
he
localized
ood
supply,
esul ed
in
some measu e
o
empo al
sepa a ion
o
eeding
in
ha
E.
gambianus
had
unlimi ed
access
o
he
ood
esou ces
o
almos
wo
hou s
p io
o
he
a i al
o
R.
aegyp iacus.
406
D.
W.
THOMAS
AND
M.
B.
FENTON
We
a e
g a e ul
o
he
Depa men
o
Na ional
Pa ks
and
Wild
Li e
Managemen o
Rhodesia
o
pe mi ing
us
o
wo k
a
he
Ins i u e
and
o
use
he
acili ies
he e.
We
a e
especially
g a e ul
o
Rowan
and
Liz
Ma in
o
hei
hospi ali y
du ing
ou
s ay,
and
we
a e
pa icula ly
indeb ed
o
Rowan
o
sha ing
his
ex ensi e
knowledge
o
eleme y
wi h
us.
We
hank
Edson
Mulilo
and
Zaccheus
Mhlangu
who
assis ed
wi h
he
ield
wo k
and
all
membe s
o
he
communi y
a
he
Ins i u e
o
hei
assis ance.
D
D.
H.
M.
Cumming
was
pa icula ly
help ul
in
all
aspec s
o
he
s udy
and
we
hank
Rudye d
Boul on
o
he
A lan ica
Ecological
Resea ch
S a ion
o
lending
us
some
equipmen .
D.
H.
M.
Cumming,
T.
H.
Fleming,
T.
H.
Kunz
and
R.
B.
Ma in
ead
he
manusc ip
and
made
help ul
sugges ions.
This
s udy
was
suppo ed
by
Na ional
Resea ch
Council
o
Canada
g an s
o
M.B.F.
and
by
he
Science
Facul y
a
Ca le on
Uni e si y.
D.
H.
M.
Cumming
kindly
p o ided
he
habi a
da a
o
Figu e
1.
REFERENCES
Ayensu,
E.
S.
(1974).
Plan
and
ba
in e ac ions
in
Wes
A ica.
Ann.
Mo.
bo .
Gcln
61:
702-727.
Bake ,
H.
G.
&
Ha is,
B.
J.
(1957).
The
pollina ion
o
Pa kia
by
ba s
and
i s
a endan
e olu iona y
p oblems.
E olu ion,
Lancas e ,
Pa.
11:
449-460.
B adbu y,
J.
W.
(1977).
Lek
ma ing
beha iou
in
he
hamme -headed
ba .
Z.
Tie psychol.
45:
225-255.
B osse ,
A.
(1966).
La
biologie
des
chi op e es.
Pa is:
Masson
e
Cie.
Cumming,
D.
H.
M.
(1975).
A
ield
s udy
o
he
ecology
and
beha iou
o
wa hog.
Mem.
na n.
Mas.
Rhod.
No.
7:
1-179.
Fen on,
M.
B.
(1975).
Obse a ions
on
he
biology
o
some
Rhodesian
ba s,
including
a
key
o
he
Chi op e a
o
Rhodesia.
Con .
Li e
Sci.
Di .
R.
On .
Mus.
No.
104:
1-27.
Jones,
C.
(1971).
The
ba s
o
Rio
Muni,
Wes
A ica.
J.
Mammal.
52:
121-140.
Jones,
C.
(1972).
Compa a i e
ecology
o
h ee
p e opid
ba s
in
Rio
Muni,
Wes
A ica.J.Zool.,Lond.
167:
353-370.
Kingdon,
J.
(1974).
Eas A ican
mammals,
an
a las
o
e olu ion
in
A ica.
2.A.
London:
Academic
P ess.
Rose ea ,
D.
R.
(1965).
The
ba s
o
Wes
A ica.
London:
B i ish
Museum
(Na u al
His o y).
Smi he s,
R.
H.
N.
(1971).
The
mammals
o
Bo swana.
Mem.
na n.
Mus.
Rhod.
No.
4:
1-340.
Smi he s,
R.
H.
N.
&
Lobao
Tello,
J.
L.
P.
(1976).
Check
lis
and
a las
o
he
mammals
o
Mozambique.
Mem.
na n.
Mus.
Rhod.
No.
8:
1-184.
J.
Zoo].,
Lond.
(1978)
186,
407-416
Obse a ions
on
he
locomo ion
o
wo
B i ish
e es ial
plana ians
(Pla yhelmin hes,
T icladida)
H.
D.
JONES
Depa men
o
Zoology,
Uni e si y
o
Manches e ,
Manches e
{Accep ed
11
Ap il
1978)
(Wi h
2
pla es
and
1
igu e
in
he
ex )
The
locomo ion
o
Mic oplana
e es is
and
o
M.
b i annicus
is
desc ibed.
Fo wa d
loco-
mo ion
is
no mally
by
means
o
cilia
which
a e
con ined
o
he
en al
su ace
o
he
animals.
M.
e es is
may
howe e
in ol e
s a iona y
pe is al ic
wa es
in
locomo ion.
In
his
case
nei he
muscula
no
cilia y
o ces
can
alone
accoun
o
he
locomo ion
and
i
is
necessa y
ha
he
wo
mechanisms
a e
combined.
Such
s a iona y
wa es
should
be
dis-
inguished
om
e og ade
and
di ec
locomo o y
wa es.
Re e sal
in
bo h
species
is
by
e og ade
muscula
wa es.
Con en s
In oduc ion
Ma e ials
and
me hods
Resul s
Mic oplana
e es is
.
Mic oplana
b i annicus
Discussion
Summa y
Re e ences
Page
407
408
408
408
410
410
415
415
In oduc ion
The
e es ial
plana ian
auna
o
G ea
B i ain
has
been
a ely
obse ed
and
is
poo ly
documen ed.
Acco ding
o
Cloudsley-Thompson
&
Sankey
(1961)
he e
a e
ou
species.
This
pape
eco ds
obse a ions
on
wo
o
hem,
Mic oplana
e es is
(M ille )
and
.
b i annicus
(Pe ci al).
Obse a ions
on
he
locomo ion
o
e es ial
plana ians
a e
ew,
he
p incipal
wo k
oeing
ha
o
Pan in
(1950).
Locomo ion
has
been
conside ed
o
be
accomplished
by
a
combina ion
o
cilia y
o ces,
cilia
being
con ined
o
he
na ow
en al
su ace,
and
muscu-
la
con ac ions
ei he
o
he
en al
su ace
o
o
he
whole
body
bu
wi h
li le
de ailed
elabo a ion
as
o
how
he
wo
may
be
combined.
Pan in
(1950)
conside s
locomo ion
o ces
o
be
p incipally
muscula
in
o igin.
Mucus
is
sec e ed
om
gland
cells
ha
a e
concen a ed
nea
he
an e io
end
and
a
slime- ail
is
le
by
he
animals.
In
some
species
locomo ion
is
supplemen ed
by
appa en
pe is al ic
wa es
o
muscula
con ac ion
which
pass
along
he
wo m
a
he
same
a e
as
he
wo m
mo es
o wa ds.
Such
wa es
hus
emain
s a iona y
ela i e
o
he
subs a um
and
wo ms
exhibi ing
hese
s a iona y
wa es
lea e
a
do ed
slime- ail.
Simila
s a iona y
wa es
a e
seen
du ing
ocomo ion
o
ce ain
land
snails
and
slugs
(Jones,
1975).
I
will
be
shown
in
his
pape
ha
407
