Long-eared Bats

Long-Ea ed
Ba s
Long-Ea ed
Ba s
Susan
M.
Swi i
Wi h
illus a ions
by
Ru h
Lindsay
T&AD
POYSER
NATURAL
HISTORY
i
Con en s
CHAPTER
4
Fo aging
beha iou
51
Me hods
o
ma king
and
acking
o aging
ba s
Ac i i y
pa e ns
Fo aging
CHAPTER
5
Rep oduc ion
68
Seasonal
cycle
Male
ep oduc ion
Female
ep oduc ion
Bi h
and
li e
size
Pos -na al
g ow h
and
de elopmen
Ene ge ics
o
ep oduc ion
CHAPTER
6
Social
o ganiza ion
and
beha iou
89
Roos ing
beha iou
The mo egula ion
in
oos s
Tempo a y
and
ansien
oos s
Nu se y
colonies
Beha iou
o
males
in
summe
Mo he —young
in e ac ion
Ma ing
beha iou
CHAPTER
7
Popula ion
biology
106
Li e
expec ancy
and
su i al
a e
Popula ion
size
Sex
a io
Fligh
dis ances
Pa asi es
P eda ion
Dayligh
lying
CHAPTER
8
Hibe na ion
119
He e o he my,
o po
and
hibe na ion
Si e
selec ion
A ousal
Con en s
VI1
Win e
ac i i y
E ec s
o
dis u bance
on
hibe na ing
ba s
CHAPTER
9
Long-ea ed
ba s
and
humans
132
Posi i e
human
in luences
Th ea s
o
ba s
Ba s
and
he
law
Public
ela ions
E ec s
o
esea ch
on
ba s
Remedial
imbe
ea men
S ee
lamps
and
u baniza ion
Ba
boxes
P ac ical
conse a ion
measu es
Appendix
1
Scien i ic
and
common
names
o
ba
species
149
Appendix
2
Ca e
o
s anded
o
inju ed
ba s
151
Appendix
3
Use ul
con ac
add esses
in
he
UK
153
Re e ences
155
Index
175
Lis
o
Colou
Pla es
Pla e
1
A
b own
long-ea ed
ba
(Pleco us
au i us)
in
ligh .
The
la ge,
e ec
ea s
can
be
seen
in
o chligh .
Pho o:
F ank
G eenaway.
Pla e
2
G ey
long-ea ed
ba
(Pleco us
aus iacus).
The
agus
shape,
da k
b own
acial
colou ing
and
sho
humbs
iden i y
his
indi idual.
Pho o:
F ank
G eenaway.
Pla e
3
A
b own
long-ea ed
ba
a
oos
wi h
ea s
in
he
' am's
ho n'
posi ion.
The
ans e se
olds
in
he
conch
close
in
a
an-like
ac ion
as
he
ea
elaxes.
Pho o:
John
Haddow.
Pla e
4
Because
long-ea ed
ba s
ly
inside
oos s,
hei
d oppings
become
sca e ed
a ound
a ics
and
may
co e
u ni u e
o
o he
possessions
s o ed
he e.
Pla e
5
P.
au i us
lying
close
o
ege a ion.
Du ing
oliage
gleaning,
b own
long-ea ed
ba s
equen ly
s op
echoloca ing
and
sea ch
o
p ey
by
passi e
lis ening.
Pho o:
F ank
G eenaway.
Pla e
6
Insec
emains
accumula e
benea h
eeding
pe ches.
Clea ly,
he
ba
ha
used
his
pe ch
had
been
hun ing
la ge
yellow
unde wing
mo hs.
Pho o:
F ank
G eenaway.
Pla e
7
Flyways
a e
consis en ly
used
by
long-ea ed
ba s
o
a oid c ossing
open
spaces
while
commu ing
be ween
oos s
and
o aging
a eas.
They
ly
close
o
ege a ion
along
landscape
ea u es
such
as
o e g own
s eams.
Pla e
8
A
g oup
o
b own
long-ea ed
ba s
in
hei
no mal
oos ing
posi ion
in
an
a ic.
They
we e
pho og aphed
om
below
and
a e
huddled
be ween
he
idge
beam
and
he
sa king,
in
he
angle
o med
by
a
a e .
Pla e
9
A
emale
b own
long-ea ed
ba
wi h
a
newbo n
in an .
Babies
a e
con inuously
a ached
o
hei
mo he 's
nipple
du ing
hei
i s
week
o
li e.
Pho o:
F ank
G eenaway.
Pla e
10
B own
long-ea ed
ba s
ha e
been
ound
o
selec
olde
houses
wi h
complex
oo
spaces.
A
colony
o
80
oos ed
in
his
Vic o ian
house
and
equen ly
mo ed
be ween
oo
compa men s.
Pla e
11
Long-ea ed
ba
oos s
a e
likely
o
be
si ua ed
in
wooded
alleys.
Well-
main ained
ishing
i e s
a e
an
impo an
ea u e
o
he
ecology
o
he
species.
Pla e
12
P.
au i us
selec
houses
which
ha e
woodland
wi hin
0.5
km.
This
oos
is
su ounded
by
ma u e
deciduous
and
coni e ous
ees.
Pla e
13
A
hibe na ing
b own
long-ea ed
ba
wi h
olded
ea s.
The
agi
hang
down,
supe icially
esembling
he
ea s
o
o he
species.
Pho o:
F ank
G eenaway.
P e ace
I
n
ecen
yea s,
a
numbe
o
books
on
he
biology,
beha iou
and
ecology
o
ba s
ha e
been
published,
in
line
wi h
inc easing
knowledge
o
he
g oup
and
de eloping
in e es
in
i .
Howe e ,
he e
ha e
been
e y
ew
in-dep h
monog aphs,
and
he ime
now
seems
igh
o
mo e
de ailed
e iews
dealing
wi h
indi idual
species.
Long-ea ed
ba s
a e
among
he
bes
known
Eu opean
species.
They
a e
dis inc i e,
easily
ecognizable
and
a ac i e,
and
he e
a e
also
aspec s
o
hei
biology
and
beha iou
which
make
hem
di e en
om
o he
ba s
and
he e o e
wo hy
o
being
conside ed sepa a ely.
I
hope
his
book
will
be
o
in e es
o
ama eu
ba
en husias s
who
wan
o
disco e
mo e
abou
long-ea ed
ba s,
as
well
as
o
p o essional
biologis s.
I
is
also
in ended
as
an
in o-
duc ion
o
he
species
o
s uden s
emba king
on
esea ch
in o
hei
biology;
I
hope
i
will
be
help ul.
Sue
Swi
May
1997
Acknowledgemen s
I
am
g a e ul
o
colleagues
in
he
Zoology
Depa men
o
he
Uni e si y
o
Abe deen,
bo h
pas
and
p esen ,
whose
ideas
and
en husiasm
I
ha e
d awn
upon
and
whose
esea ch
on
long-ea ed
ba s
has
o med
he
basis
o
much
o
his
book.
In
pa icula
I
hank
P o esso
Paul
Racey,
who
was
la gely
esponsible
o
my
ini ial
in e es
in
ba s
and
who
supe ised
and
encou aged
all
my
ea ly
wo k;
he
has
also
allowed
me
unlimi ed
access
o
his
collec ion
o
li e a u e,
o
which
many
hanks
a e
due.
In
addi ion,
I
am
indeb ed
o
Abigail
En wis le
o
access
o
he
unpublished
PhD
hesis.
Tony
Hu son,
Jens
Rydell,
Abigail
En wis le,
Paul
Racey
and
John
Speakman
ead
and
imp o ed
ea ly
d a s
o
a ious
chap e s
and
de o ed
ime
and
e o
o
inding
e e ences
which
I
had
missed
—
I
am
mos
g a e ul
o
hem
all.
I
also
hank
he
lib a y
s a
a
he
Uni e si y
o
Abe deen
o
help
in
acing some
o
he
li e a u e.
Du ing
wen y
yea s
o
ield
wo k,
I
ha e
ecei ed
hospi ali y
and
coope a ion
om
many
oos
owne s;
I
should
like
o
ake
his
oppo uni y
o
hank
hem
all
and
o
acknow-
ledge
he
in aluable
con ibu ion
which
oos
owne s
make
o
ba
conse a ion
h oughou
Eu ope.
Finally,
I
am
g a e ul
o
my
amily,
John,
Imogen
and
Ma cus.
Besides
adap ing
o
cope
wi h
my
noc u nal
way
o
li e
e e y
summe ,
hey
ha e
ole a ed
an
e en
wo se
han
usual
s anda d
o
housekeeping
du ing
he
mon hs
I
was
w i ing
and
ha e
enabled
me
(mo e
o
less!)
o
main ain
a
sense
o
humou
h oughou .
CHAPTER
1
Who's
Who
—
An
In oduc ion
o
Long-Ea ed
Ba s
T
HE
aim
o
his
book
is
o
desc ibe
he
biology,
ecology
and
beha iou
o
Eu opean
ba
species
o
he
genus
Pleco us.
Long-ea ed
ba s a e
among
he
mos
dis inc i e
and
bes
known
o
ou
ba
auna
and
a e
immedia ely
ecognizable
by
hei
eno mous
ea s
which
a e
abou
h ee-qua e s
o
he
leng h
o
he
es
o
he
ba .
Wi h
hese
huge
ea s
and
hei
la ge
eyes
and
delica e
aces,
Pleco us
ba s a e
physically
e y
a ac i e
and
pho og aphs
o
hem
ha e
appea ed
on
he
co e s
o
se e al
ecen
ba
books
aimed
a
he
gene al
public.
They
a e
also
among
he Eu opean
species
mos
closely
associa ed
wi h
buildings,
and
as
such
a e
likely
o
be
encoun e ed
by
membe s
o
he
public.
Because
hey
a e
so
a ac i e,
hey
ha e
become
seen
as
an
impo an
asse
in
he
all-impo an
'Ba s
Need
F iends'
campaign
o
imp o e
he
public
image
o
all
ba s.
They
ha e
been
ea u ed
on
a
numbe
o
logos
in
ecen
yea s,
including
hose
o
he Eu opean
Ba
Resea ch
O ganiza ion
and
he
Ba
Conse a ion
T us .
Because
hey
equen ly
oos
in
houses
and
because
o
hei
habi
o
using
eeding
pe ches
and
he e o e
lea ing
in e es ing
heaps
o
insec
emains
o
humans
o
ind,
hey
a e
also
ba s
which
ha e
long
been
o
in e es
o
ama eu
na u alis s.
Knowledge
o
hei
biology
and
beha iou
has
ad anced
apidly
o e
he
las
wo
decades;
hei
o aging
me hods
a e
di e en
om
hose
o
o he
Eu opean
species,
and
in
a
numbe
o
o he
ways
hey
a e
also
wo hy
o
being conside ed
sepa a ely.
All
ba s
a e
no
alike,
jus
as
all
oden s
a e
no
alike,
and
I
hope
he
ollowing
chap e s
will
begin
o
show
how
di -
e en
and
how
ascina ing
hese
ba s
a e.
Many
ba s
wo ldwide
ha e
la ge
ea s,
and
he
names
'big-ea ed'
and
'long-ea ed'
ha e
been
widely
applied
o
species
in
no
ewe
han
nine
mic ochi op e an
amilies.
Howe e ,
he
gene a
Pleco us,
Co yno hinus,
Idionyc e is
and
Eude ma
in
he
amily
Vespe ilionidae
a e
peculia
in
ha
hey
ha e
he
abili y
o
old
hei
ea s
while
hey
a e
a
es .
In
e ec ,
he
ea s
a e
'de la ed'
(Hill
and
Smi h,
1984)
by
he
closing
o
special
al es
in
blood
essels
which
en e
he
ea
conch,
hus
allowing
he
ea
o
collapse.
When
he
ba
becomes
ac i e,
he
ascula
al es
a e
opened
and
he
in ushing
blood
causes
he
ea
o
become
e ec .
The
genus
Pleco us
con ains
wo
Palea c ic
species
which
a e
widesp ead,
a
hi d
con ined
o
he
Cana y
Islands
and
a
ou h
in
Taiwan.
The e
a e
also
h ee
No h
Ame ican
species,
which
a e
now
e-classi ied
as
Co yno hinus
(see
Close
Rela ions,
below).
The
b own
long-ea ed
ba ,
Pleco us
au i us,
L.
1758,
occu s
widely
in
Eu ope
and
is
p ob-
ably
he
second
commones
B i ish
ba a e
he
pipis elle,
Pipis ellus
pipis ellus
(Mi chell-Jones,
1990).
I
is
a
small
o
medium-sized
ba
o
delica e
build,
weighing
1

2
Who's
Who
—
An
In oduc ion
o
Long-Ea ed
Ba s
5-10
g
and
wi h
ea s
a
leas
25
mm
long
joined
a
hei
bases
(S ebbings,
1986).
These
may
be
cu led,
esembling
ams'
ho ns
when
he
ba
is
a
es .
The
agus
(inne
ea )
is
slen-
de
and
he
ace
colou
is
usually
b own
o
pink.
The
do sal
pelage
is
b own,
wi h
da k
and
ligh
bands
along
he
leng h
o
he
hai s.
The
g ey
long-ea ed
ba ,
Pleco us
aus iacus,
Fische ,
1829,
is
a e
in
B i ain
bu
is
wide-
sp ead
in
Eu ope,
pa icula ly
in
Medi e anean
a eas.
I
is
e y
simila
o
P.
au i us,
bu
is
o e all
sligh ly
la ge
(7-14
g)
and
i s
coa
colou
g eye ,
wi h
hai s
e enly
colou ed
along
hei
whole
leng h,
al hough
he e
is
conside able
o e lap
in
bo h
size
and
coa
colou .
I s
ace
is
g ey
o
black
and
i s
humbs
a e
sho
and
hick.
The
wo
species
a e
e y
di icul
o
dis inguish,
especially
when
dealing
wi h
li ing
specimens,
and
his
has
led
o
conside able
con usion
in
a
numbe
o
eco ds
o
hei
dis ibu ion,
especially
in
pa s
o
hei
anges
whe e
hey
a e
sympa ic.
A
hi d
species,
Pleco us
ene i ae
(Ba e -Hamil on,
1907),
is
ound
only
in
he
Cana y
Islands,
whe e
isola ion
has
led
o
he
e olu ion
o
his
endemic
species
(Boye
e
ai,
1990).
Al hough
i
is
la ge
han
P.
au i us
and
al hough,
om
geog aphic
dis ibu ion,
i
has
mo e
in
common
wi h
he
sou he n
species
P.
aus iacus,
Ibanez
and
Fe nandez
(1986)
consid-
e ed
i
o
be
mo e
closely
ela ed
o
P.
au i us
han
o
P.
aus iacus.
Li le
is
known
o
he
ecology
o
beha iou
o
he
Tene i e
long-ea ed
ba .
Long-ea ed
ba s
in
Taiwan,
o me ly
conside ed
o
be
P.
au i us
(S ebbings
and
G i i hs,
1986),
a e
now
ecognized
as
a
sepa-
a e
species,
Pleco us
ai anus
(Yoshiyuki,
1991).
This au ho
also
ad oca ed
sepa a e
species
s a us
o
P.
au i us
in
Nepal
and
India,
bu
his
has
no
been
ecognized
by
o he
au ho i ies.
HISTORY
AND
ORIGIN
OF
THE
SPECIES
The
di icul y
in
dis inguishing
Pleco us
species
has,
in
he
pas ,
led
o
con usion.
Fo
many
yea s,
Eu ope
was
conside ed
o
ha e
only
one
species,
P.
au i us;
in
1907,
P.
ene i ae
was
desc ibed,
and
i
was
no
un il
ela i ely
ecen ly
ha
he
exis ence
o
P.
aus iacus
as
a
species
a he
han
a
a ie y
o
P.
au i us
was
ecognized.
Jenyns
(1829)
desc ibed
wo
species
o
long-ea ed
ba s
in
B i ain,
calling
he
la ge ,
g eye
one
P.
au i us
and
he
smalle ,
b owne
one
P.
b e imanus.
As
Co be
(1964)
poin ed
ou ,
i
hese
did,
indeed,
cons i u e
wo
species,
hen
Jenyns
had
hem
he
o he
way
ound
o
he
way
in
which
we
now
accep
he
si ua ion!
Bell
(1874)
ejec ed
P.
b e imanus
as
being
he
young
o
P.
au i us.
Simila ly,
in
Eu ope,
wo
o ms
we e
ecognized
(Kuzyakin,
1944;
Topal,
1958)
bu
conside ed
o
be
subspecies,
P.
au i us au i us
and
P.
a.
wa di,
Thomas.
I
was
no
un il
1959
ha
he
exis-
ence
o
wo
good
species,
occu ing
sympa ically
in
I aly,
was
shown
(Lanza,
1959).
Lanza
desc ibed
he
smalle ,
b owne
species
as
P.
au i us
and
he
la ge ,
g eye
one
as
P.
wa di.
A e
his,
he
p esence
o
wo
sympa ic
species
was
shown
in
Aus ia
(Baue ,
1960),
Czechoslo akia
(Hanak,
1962)
and
he
Ne he lands
(Wijngaa den,
1962).
Baue
(1960)
named
he
g ey
species
P.
aus iacus,
which
was
he
name
o iginally
gi en
by
Fische
in
1829
o
a
a ie y
o
P.
au i us
om
Aus ia.
Co be
(1964)
e-examined
specimens
o
P.
au i us
in
he
B i ish
Museum
(Na u al
His o y)
and
disco e ed
among
hem
se e al
indi iduals
o
P.
aus iacus.
One
o
hese
had
been
collec ed
in
Hampshi e
be ween
1873
and
1878,
one
came
om
Je sey
in
he
Channel
Islands
and
he
es
we e
om
mainland
Eu ope.
The
p esence
o
P.
aus iacus
in
Dis ibu ion
3
B i ain
was
hus
es ablished.
Co be
(1964)
concluded
ha
i
was
ei he
e y
local
in
his
coun y
o
ha
i
was
much
less
abundan
han
P.
au i us-,
bo h
conclusions
ha e
since
been
shown
o
be
co ec .
Sho ly
a e
his,
S ebbings
(1966)
iden i ied
bo h
species
in
a
colony
in
Do se
which
he
had
been
s udying
o
some
ime.
Mixed
colonies,
whe e
he
wo
species
appa en ly
li e
in
close
p oximi y
o
each
o he ,
ha e
subsequen ly
been
epo ed
widely
in
hei
common
ange.
E en
when
he
p esence
o
bo h
is
suspec ed,
iden i ica ion
can
be
e y
di icul .
F om
he
abo e,
i
is
ob ious
ha
o
a emp
o
a ibu e
eco ds
o
Pleco us
ba s
om
be o e
1959
o
one
species
o
he
o he
would
be
un eliable
and
so,
p io
o
his
da e,
eco ds
a e
e e ed
o
as
Pleco us
spp.
unless
hey
we e
om
places
whe e
only
one
species
occu s.
Whe e
iden i ica ion
canno
be
made
wi h
ce ain y
(e.g.
in
many
hibe na ion
eco ds
o
in
o he
cases
whe e
ba s
could
no
be
handled),
e en
hose eco ded
since
1959
a e,
in
many
cases,
iden i ied
only
o
genus.
Ho acek
(1975)
a ibu ed
many
o
he
di e ences
in
he
ecology
o
P.
au i us
and
P.
aus iacus,
such
as
in
die
and
o aging
(see
Chap e s
3
and
4),
oos ing
beha iou
and
ma -
ing
sys ems
(Chap e
6)
and
choice
o
hibe na ion
si es
(Chap e
8),
o
di e ences
in
he
his o y
o
he
wo
species
in
Eu ope.
While
he
ecology
o
P.
au i us
is
ha
o
an
old
bu
s ill
adap able
species,
ha
o
P.
aus iacus
indica es
ha
i
is
a
new
species
in
he
a ea
and
one
which
has
sp ead
only
ecen ly.
P.
au i us
is
known
om
he
ossil
eco d
o
ha e
been
p esen
in
Eu ope
om
he
Pliocene
( om
12
million
o
2—3
million
yea s
ago)
and
o
ha e
been
common
in
he
Pleis ocene,
up
o
2
million
yea s
ago
(Ho acek,
1975;
Sese
and
Ruiz-
Bus os,
1992).
The
species
hus
appea s
o
ha e
i s
o igins
in
Eu ope.
P.
aus iacus,
on
he
o he
hand,
is
Asian
in
o igin
and
is
a
ema kably
new
species
in
Eu ope
(Ho acek,
1975),
sp eading
h ough
he
con inen
in
his o ical
ime.
I
appea s
o
ely
hea ily
on
human
dwellings
as
summe
oos
si es
and
on
a i icial
s uc u es
such
as
cella s
as
hibe nacula,
and
i
is
possible
ha
i
only
sp ead
in
Eu ope
as
such
si es
became
a ailable.
DISTRIBUTION
Bo h
P.
au i us
and
P.
aus iacus
a e
widesp ead
in
he
Palea c ic
Region.
P.
au i us
occu s
eas wa d
as
a
as
he
U al
and
Caucasus
moun ains.
In
Asia,
i
is
ound
in
Mongolia
and
no h-eas
China
(Zeng
and
Wang,
1989),
Sakhalin
and
Japan
(Co be
and
Hill,
1991).
I
also
occu s
in
Sou h
Ko ea
and
in
isola ed
pocke s
in
cen al
China,
as
well
as
in
pa s
o
India
and
Nepal
(Co be
and
Hill,
1991).
P.
aus iacus
is
dis ibu ed
in
sou he n Eu ope
and
no h
A ica,
eas wa ds
o
Mongolia
and
wes e n
China
and
sou hwa ds
o
Senegal
(Nowak,
1991)
and
E hiopia
(La gen
e
al.,
1974).
In
Eu ope,
P.
aus iacus
is
he
com-
mone
species
in
sou he n
coun ies
whe e
he
clima e
is
Medi e anean
and
ela i ely
s able.
The
wo
species
occu
oge he
in
cen al
Eu ope
o
app oxima ely
53°N,
and
P.
au i us
occu s
on
i s
own
a
highe
la i udes
(up
o
64°N),
whe e
he
clima e
is
coole
and
mo e
changeable
(S ebbings,
1970).
Figu es
1.1
and
1.2
show
he
dis ibu ion
o
P.
au i us
and
P.
aus iacus
espec i ely.
P.
au i us
is
common
and
widesp ead
in
B i ain
(Figu e
1.3).
I
occu s
e e ywhe e
excep
in
exposed
moun ainous
egions
in
no h
and
no h-wes
Sco land
(Swi ,
1991;
A nold,
1993).
I
is
ound
in
he
Inne
Heb ides
and
has
been
eco ded
on
O kney
(Boo h
and
Boo h,
1994),
bu
is
absen
om
he
ou e
islands
and
She land.
I s
absence
om
hese
c-
K
s
Q
O
E
6
Who's
Who
—
An
In oduc ion
o
Long-Ea ed
Ba s
a eas
may
well
be
due
o
hei
lack
o
ees,
since
P.
au i us
is
s ongly
associa ed
wi h
wood-
land
(Swi
and
Racey,
1983).
I
is
common
h oughou
I eland
(Mo a ,
1938;
O'Sulli an,
1994),
whe e
i
occu s
in
la ge
numbe s
in
many
a eas
(O'Go man
and
Fai ley,
1965).
In
B i ain,
P.
aus iacus
is
a e
and
is
con ined
o
he
ex eme
sou h
o
he
coun y
(Figu e
1.4);
i
has
been
eco ded
om
Some se ,
Do se ,
De on,
Hampshi e,
he
Isle
o
Wigh
and
he
Channel
Islands
(A nold,
1993).
The e
a e
se e al
eco ds
om
he
ex eme
wes
o
Sussex
and,
mo e
ecen ly,
also
om
B igh on
(Hu son,
1991)
and
Chiches e
(Hu son,
1996).
I
is
epo ed
o
be
he
second
commones
ba
in
Je sey
a e
he
pipis elle
(D.
La oley,
pe s.
comm.)
and
i
has
been
eco ded
on
all
he
Channel
Islands
excep
He m.
I
is
known
o
b eed
on
Je sey
and
Gue nsey
and,
possibly,
also
on
Alde ney;
he
Channel
Islands a e
hough
o
con ain
abou
h ee-qua e s
o
he
B i ish
popula ion
o
his
species
(D.
La oley,
pe s.
comm.).
P.
aus iacus
is
absen
om
Scandina ia,
bu
P.
au i us
is
common
in
Denma k
and
occu s
h oughou
he
coun y
(Baagoe,
1980-81).
I
is
also
common
in
cen al
and
sou h-
e n
Sweden
(Ge ell,
1980-81)
and
occu s
as
a
no h
as
la i ude
63°
o
64°N
(Ahlen
and
Ge ell,
1989;
Schobe
and G immbe ge ,
1989).
I
is
p esen
o
simila
la i ude
in
No way,
Dis ibu ion
7
FIG
1.4
Dis ibu ion
o
P.
aus iacus
in
he
B i ish
Isles.
in
sou he n
Finland
and
in
Es onia.
In
Russia,
i
has
been
eco ded
as
a
no h
as
60°25'N
(Re in-Yu
and
Boesko o ,
1989).
Bo h
species
occu
h oughou
F ance,
Swi ze land
and
Aus ia.
On
he
Aus ian-Czech
bo de ,
high
numbe s
o
bo h
we e
ne ed
in
wooded
a eas
(Gaisle
e
al.,
1996)
and,
also
in
Aus ia,
bo h
we e
ound
a
heigh s
o
350—1500
m
abo e
sea
le el
(Spi zenbe ge ,
1993).
P.
aus iacus
we e
ound
mainly
in
submon ane
a eas
and
P.
au i us
mainly
in
mon-
ane
a eas.
Bo h
also
occu
in
he
Ne he lands
and
Belgium.
Joo is
(1980)
epo ed
ha
P.
aus iacus
was
ai ly
common
in
he
low-lying
dis ic s
o
Belgium,
whe e
i
appea ed
o
p e e
mo e
open
habi a s
han
he
woodland
a eas
inhabi ed
by
P.
au i us.
In
he
Ne he lands,
P.
aus iacus
is
uncommon
and
is
mos ly
con ined
o
he
sou he n
hal
o
he
coun y
(Daan,
1980;
Glas,
1982).
P.
au i us
was
o me ly
common,
bu
appea s
o
ha e
su e ed
a
sha p
d op
in
numbe s
ecen ly
(Daan,
1980).
The
wo
species
occu
in
he
Czech
and
Slo ak
Republics
(Gaisle
e
al.,
1980—81).
Numbe s
o
bo h
a y
bu
a e
locally
high
in
o es ed
a eas
(Gaisle
e
al.,
1996)
and
in
a eas
whe e
sui able
hibe nacula
a e
ound
(Weidinge ,
1994).
In
Poland,
P.
aus iacus
eaches
he
no he n
limi
o
i s
8
Who's
Who
—
An
In oduc ion
o
Long-Ea ed
Ba s
dis ibu ion
a
abou
53°N
(i.e.
sligh ly
u he
no h han
in
B i ain,
whe e
i
is
no
ound
abo e
51°N)
and
is
hus
p esen
in
he
sou he n
wo- hi ds
o
he
coun y
(Rup ech ,
1971).
In
cen al
and
wes
Poland
i
is
gene ally
a
lowland
species,
bu
in
he
eas
i
ex ends
o
oo hill
egions.
P.
au i us
occu s
h oughou
he
coun y,
and
he
wo
species
a e
hus
sympa ic
below
53°N.
Simila ly
in
Ge many,
bo h
species
a e
ound
oge he
as
a
no h
as
a ound
Be lin
a
52°30'N
(Haensel
and
Na e,
1993),
bu
P.
aus iacus
has
no
been
eco ded
u he
no h.
Bo h
species
occu in
Bulga ia
and
Romania
(S ebbings
and
G i i hs,
1986).
Fu he
sou h
in
Eu ope,
P.
aus iacus
is
he
commone
and
mo e
widesp ead
o
he
wo
species.
Bo h
occu
in
Po ugal,
bu
P.
au i us
is
a e
and
was
only
ecen ly
eco ded
o
he
i s
ime
(Palmei im,
1990).
P.
aus iacus
occu s
h oughou
Spain
and
in
he
Balea ic
Islands
( om
which
P.
au i us
is
absen )
and
li es
a
a
ange
o
al i udes
om
sea
le el
o
1600
m,
al hough
i
is
only
in
he
sou h
ha
i
inhabi s
moun ain
a eas
(de
Paz,
1984).
An
examina ion
o
museum
specimens
o
Pleco us
collec ed
na ionwide
e ealed
ha
77%
we e
P.
aus iacus
and
23%
we e
P.
au i us
(de
Paz,
1984).
P.
au i us
appea s
o
be
widely
dis-
ibu ed
in
he
no he n
hal
o
Spain,
as
a
sou h
as
Sie a
de
Guada ama
and
G edos
in
he
p o ince
o
Mad id
(de
Paz,
1984;
Benzal,
1991).
I
occu s
a
all
al i udes
in
he
no h,
bu
only
highe
han
1000
m
abo e
sea
le el
u he
sou h.
I
is
commones
in
highland
egions
such
as
he
Py enees,
Can ab ian
and
Galician
moun ains,
whe e
i
gene ally
li es
in
ei he
deciduous
o
Pinus
o es
(de
Paz,
1984).
In
he
Py enees,
P.
au i us
was
epo ed
o
be
he
commones
ba
species
ne ed
a
al i udes
o
2000-2500
m
(Be and,
1992).
I
does
no
appea
o
be
p esen
in
he
sou he n
hal
o
Spain.
I
occu s
in
I aly
as
a
sou h
as
app oxima ely
43°N
(C uci i,
1989)
and
is
absen
om
Co sica,
Sa dinia
and
Sicily.
P.
aus iacus
occu s
h oughou
mos
o
mainland
I aly
excep
he
sou hwes
bu
is
simila ly
absen
om
Co sica,
Sa dinia
and
Sicily
(Ve nie ,
1987).
P.
aus iacus
is
he
commone
o
he
wo
species
in
he
o me
Yugosla ia
and
is
widely
dis ibu ed
he e
(K ys u ek,
1980).
Two
popula ions
ha e
been
iden i ied
and
hese
di e
mainly
in
body
size
(Dulic,
1980).
The
la ge
specimens,
o
simila
size
o
hose
o
his
species
elsewhe e
in
Eu ope,
a e
ound
in
no he n and
sou h-eas e n
a eas,
while
smalle
ones
occu
along
he
Ad ia ic
coas
and
on
adjacen
islands.
Popula ions
in
cen al
C oa ia
and
Bosnia
con ain
specimens
wi h
cha ac e is ics
in e media e
be ween
he
wo.
Dulic
(1980)
p oposed
naming
he
smalle
specimens
as
a
subspecies,
P.
aus iacus
kolomba o ici.
Dis ibu ion
o
P.
au i us
in
he
a ea
is
much
mo e
limi ed;
i
occu s
in
C oa ia,
Bosnia
and,
uncommonly,
in
Slo enia
(K ys u ek,
1980).
P.
aus iacus
is
also
much mo e
p e alen
han
P.
au i us
in
eas e n
Medi e anean
and
Middle
Eas e n
coun ies
(de
Blase,
1980).
Specimens
o
P.
aus iacus
ha e
been
collec ed
in
Egyp
(Madkou ,
1989),
Jo dan
(Re in-Yu
and
Boesko o ,
1989;
Qumsiyeh
e
al.,
1992),
Alge ia
(Gaisle ,
1983—4)
and
Mal a
(Bo g
e
al.,
1990).
The
sou he n
limi
o
he
species
appea s
o
be
eached
in
he
Cape
Ve de
Islands
(Azza oli-Pucce i
and
Za a,
1988),
an
isola ed
island
g oup
in
he
A lan ic
Ocean
a
15°N.
I
is
ela i ely
common
in
Tu key,
G eece
and
no he n
I an
(S eine
and
Gaisle ,
1994)
and
in
Cyp us
(Boye
e
al.,
1990).
Al hough
P.
au i us
is
a e
in
his
a ea,
i
does
occu
in
moun ainous,
wooded
habi a s.
Two
specimens
we e
mis -ne ed
in
Tu key
a
600
m
abo e
sea
le el
and
one
was
caugh
a
Assalem
in
no he n
I an
(S eine
and
Gaisle ,
1994).
The
species
is
also
said
o
be
egula ly
dis ibu ed
in
he
highe
moun ain
o es s
o
G eece
(Hel e sen
and
Weid,
1990).
Thus,
while
P.
aus iacus
p e ails
in
sou he n
Eu ope
and
has
a
well-de ined
no he n
limi
o
i s
Close
Rela ions
9
ange,
p obably
delimi ed
by
i s
inabili y
o
su i e
he
cool,
changeable
clima e
a
high
la i ude,
P.
au i us
is
able
o
ex end
i s
ange
sou hwa ds
by
exploi ing
moun ainous
a eas
and
aking
ad an age
o
he
coole
clima e
in
such
a eas.
I
is
adap ed
o
li ing
in
cool,
em-
pe a e
clima es
and
so
i s
ange
ex ends
well
in o
no he n
Eu ope.
CLOSE
RELATIONS
The
axonomy
o
long-ea ed
ba s
and
hei
ela i es
has
long
been
a
subjec
o
con usion
and
changing
opinions.
Fi e
gene a
cons i u e
he
espe ilionid
ba s
known
as
he
pleco-
ine
g oup;
hey
a e
Ba bas ella,
Pleco us,
Co yno hinus,
Idionyc e is
and
Eude ma
(Koopman
and
Jones,
1970).
O
hese,
only
Ba bas ella
seems
o
ha e
escaped
he
con u-
sion.
A
genus
o
wo
Palea c ic
species,
hese
ba s
ha e
b oad,
iangula
ea s
which
a e
joined
a
he
base
and
which
do
no
old
when
he
ba
is
a
es
(S ebbings,
1991).
Pleco us
was
o iginally
applied
only
o
Palea c ic
long-ea ed
ba s,
while
h ee
e y
simila
species
in
No h
Ame ica,
known
as
big-ea ed
o
lump-nosed
ba s,
we e
classi ied
unde
he
genus
Co yno hinus.
Following
Handley's
(1959)
e ision
o
he
g oup,
howe e ,
i
was
decided
by
mos axonomis s
ha
Pleco us
and
Co yno hinus
wes
so
simila
ha
hey
should
be
con-
side ed
as
a
single
genus.
Thus,
he
Ame ican
species
we e
placed
in
a
subgenus,
Co yno hinus,
wi hin
he
genus
Pleco us
and
became
gene ally
known
as
Pleco us
ownsendii,
Pleco us
a inesquii
and
Pleco us
mexicanus.
The
single
species
o
Idionyc e is,
I.
phyllo is
was,
a
one
ime,
classi ied
as
Pleco us
phyllo is
(Ba bou
and
Da is,
1969)
bu
la e
placed
in
i s
own
genus
(Co be
and
Hill,
1991;
Nowak,
1991).
The
genus
Eude ma
con ains
one
species,
E.
macula um.
Recen ly,
phylogene ic
echniques
ha e
been
used
o
e-e alua e
ela ionships
among
pleco ine
ba s.
Using
his
me hod,
F os
and
Timm
(1992)
a gued
ha
Idionyc e is
and
Eude ma
we e
a
mo e
simila
han
we e
Idionyc e is
and
Pleco us
and
hey
should
be
syn-
onomized.
They
also
p oposed
ha
Co yno hinus
should
be
emo ed
om
he
synonym
o
Pleco us,
al hough
he
ela ionships
wi hin
Co yno hinus
and
Pleco us
we e
no
ully
esol ed.
Tumlison
and
Douglas
(1992)
in es iga ed
phylogene ic
ela ionships
among
he
g oup,
using
32
cha ac e is ics
o
he
skin
and
skull.
Cladis ic
analysis
yielded
one
mos -
pa simonious
ee.
The
cladog am
indica ed
ha
each
o
he
axa
should
be
ega ded
as
a
genus,
which
suppo ed
he
con en ion ha
Idionyc e is
is
a
dis inc
genus
(mo e
closely
ela ed
o
Eude ma
han
o
he
o he s
bu
s ill
dis inc )
and
a gued
agains
he
subgene ic
designa ion
o
Co yno hinus.
The
au ho s
he e o e
ad oca ed
ele a ing
Co yno hinus
o
ull
gene ic
s a us
and
limi ing
Pleco us
o
Palea c ic
species.
This
conclusion
is
cong uen
wi h
ha
o
Bogdanowicz
and
Owen (1996),
who
analysed
ela ionships
among
pleco ine
species
by
means
o
a
s udy
o
mo phome ic
di e gence
o
skull
cha ac e is ics.
The
Ame ican
pleco ine
ba s
hus
o m
he
ollowing
i e
species,
assigned
o
h ee
gene a.
Co yno hinus
ownsendii.
Townsend's,
o
wes e n,
big-ea ed
ba
is
widely
dis ibu ed
and
i s
ange
includes
mos
o
wes e n
No h
Ame ica
om
B i ish
Columbia
in
Canada,
sou h
o
Mexico
and
eas wa ds
h ough
cen al
USA
om
no he n
Texas
o
wes e n
Vi ginia
(Ross,
1967).
Colonies,
including
nu se y
colonies
o
up
o
1000
emales
and
young,
li e
p ima ily
in
ca es
(Humph ey
and
Kunz,
1976),
al hough
a ics
and
ba ns
a e also
used
(Ross,
1967).
Besides
he
main
popula ion
(known
as
C.
ownsendii
ownsendii),
wo
o he
subspecies
a e
ecognized
and
bo h
a e
cu en ly
conside ed
o
be
endange ed:
22
Recogni ion
and
Design
Fea u es
EARS
The
ea s
o
Pleco us
ba s
ha e
been
a
sou ce
o
in e es
o
na u alis s
o
many
yea s,
and
he e
a e
equen
desc ip ions
o
hem
in
ba
li e a u e.
Among
hese
is
one
by
Bingley
(1809),
who
kep
a
small
g oup
o
long-ea ed
ba s
and
obse ed
ha
hei
hea ing
was
'acu e
and
delica e'.
He
sugges ed
ha
hea ing
migh
well
be
he
sou ce
o
he
dex e i y
wi h
which
e en
blinded
ba s
we e
able
o
a oid
objec s
-
he
so-called
'six h
sense'
o
ba s
desc ibed
by
Spallanzani
(1784).
Bingley
(1809)
also
desc ibed
he
olding
o
he
ba s'
ea s
du ing
sleep.
He
called
he
agus
he
'inne
ea '
and
deduced
ha
i s
pu pose
was
o
keep
ou
noxious
insec s
and
o
p e en
any
o he
ex aneous
ma e
om
en e ing
he
ba 's
head
du ing
sleep.
By
olding
he
ea s
and
ucking
hem
unde
he
wings,
he
ba
hus
p e en ed
in ude s
om
en e ing.
Al hough
his
sugges ed
unc ion
o
he
agus
has
been
supe -
seded,
he
idea
ha
by
olding
he
pinnae
du ing
sleep
he
ba
is
p o ec ing
hem
om
in a-
sion
by
dus
o
pa asi es
is
s ill
conside ed
o
be
alid.
The
ea
conch
is
hin
and
memb anous,
wi h
22—24
ans e se
olds
in
bo h
species
(Schobe
and
G immbe ge ,
1989).
These
close
in
an
ac ion
simila
o
a
an
as
he
ea
is
olded.
The
inne
(an e io )
bo de
is
sligh ly
cu ed
and
is
b oadened,
wi h
a
inge
o
ine
hai s.
The
agus
is
long
and
lance -shaped
and
poin s
o wa ds
e en
when
he
ea
is
olded.
The
medial
lobules
a e
small
lobes
which
p ojec
la e ally
a
a
poin
jus
abo e
whe e
he
ea s
mee
(Haywa d
and
Teagle,
1961).
When
he
ea s
a e
e ec ,
he
lobules
ouch
each
o he .
They
a e
hough
possibly
o
be
in ol ed
in
he
olding
and
un olding o
he
pinnae
(Howa d,
1995),
an
ac ion
con olled
by
blood
lowing
in
and
ou
o
he
ea
h ough
spe-
cial
al es
(see
Chap e
1).
Haywa d
and
Teagle
(1961)
obse ed
he
medial
lobules
o
a
cap i e
b own
long-ea ed
ba
and
no ed ha
when
he
ba
was
o pid
he
lobules
we e
pale
lesh-colou ed,
conside ably
swollen
and
exuded
a
colou less
liquid.
When
i
was
ac i e
in
summe
hey
we e
lesh-pink
in
colou ,
hickened
along
he
ou e
ma gins
and
cu ed
sligh ly
inwa ds.
In
his
ba ,
which
was
in e mi en ly
ac i e
in
win e ,
he
lobules
we e
sh i elled
and
appea ed
da k
ed
in
colou
du ing
ac i e
pe iods.
The
change
om
swollen
o
sh i elled
ook
place
qui e
apidly
(o e
a
ew
hou s)
and
occu ed
be o e
he
ba
changed
om
being
o pid
o
ac i e
and
back.
Howa d
(1995)
made
a
s udy
o
he
medial
lobules;
he
obse ed
no
changes
in
hei
appea ance
and
ne e
saw
any
da k
ed
colou a-
ion.
Thei
p ecise
unc ion
emains
obscu e.
Folding
he
Pinnae
Long-ea ed
ba s
ly
wi h
hei
ea s
ully
e ec
—
he
o wa d-poin ing
pinnae
can
clea ly
be
seen
on
hose
lying
in
o chligh
and
on
ba s
caugh
in
ligh .
The
ea s
a e
e ec ed
sho ly
be o e
aking
ligh
(Schobe
and
G immbe ge ,
1989).
My
own
obse a ions
in
summe
oos s
indica e
ha ,
in
long-ea ed
ba s
which
a e
no
o pid,
e ec ion
o
he
ea s
also
occu s
as
a
eac ion
o
dis u bance.
As
he
ea s
become
e ec ,
he
ba
u ns
i s
head
as
i
lis ening
in en ly
and
hen
akes
ligh .
The
no mal
posi ion
o
he
ea s
in
oos s
whe e
ba s
a e
no
o pid
is
he
am's
ho n
posi-
ion
(Figu e
2.1b),
in
which
hey
a e
laccid
and
cu ed
backwa ds
bu
a e
no
olded.
As
he
ea s
elax,
he
ou e
edges
c inkle
up
like
a
conce ina
and
he
inne
edges
cu e
o e
and ou wa ds
(C anb ook,
1963a).
Wi h
he
ea s
in
his
posi ion,
ba s
g oom
hemsel es
Ea s
23
and
also
consume
p ey
a
eeding
pe ches.
Cap i e
long-ea ed
ba s
can
be
obse ed
mo ing
abou
hei
cage,
eeding
and
g ooming
wi h
am's
ho n
ea s.
Fully
olded
ea s
mos
commonly
occu in
hibe na ing
Pleco us,
al hough
hey
can
also
be
ound
on
o pid
ba s
in
summe
oos s
(Speakman,
1988).
The
ea s
a e
olded
longi udinally,
s e ched
ou
o
almos
hei
ull
leng h
and
ucked
unde
he
wings
pa allel
wi h
he
o e-
a ms
(Figu e
2.
lc),
wi h
he
agi
s ill
poin ing
downwa ds
and
gi ing
he
imp ession
o
long,
poin ed
ea s.
C anb ook
(1963a)
obse ed
ha
long-ea ed
ba s
did
no
old
hei
ea s
in
his
way
i
hey
we e
asleep
in
a
wa m
oom,
and
concluded
ha
he
eason
o
he
olding
was
p obably
connec ed
wi h
he mo egula ion.
Because
mos
eco ds
o
olded
ea s ha e
con-
ce ned
hibe na ing
ba s,
i
has
been
assumed
o e
a
numbe
o
yea s
ha
ea
olding
does
ha e
a
he mo egula o y
unc ion.
Many
ield
obse a ions
ha e
indica ed
ha
long-ea ed
ba s
wi h
olded
pinnae
a e
o pid
while hose
wi h
e ec
pinnae
ha e
high
body
empe a-
u es
and
a e
ac i e
(e.g.
S ebbings,
1986).
In
o de
o
cla i y
he
posi ion
and
o
in es iga e
whe he
he
deg ee
o
ea
olding
could
be
used
as
an
indica o
o
he
he mo egula o y
s a e
o
ba s
wi hou
ha ing
physically
o
handle
hem,
Speakman
(1988)
conduc ed
a
s udy
o
assess
he
ela ionship
be ween
pinna
posi ion
and
he mo egula o y
s a e
du ing
summe
and
au umn.
He
ound
ha
he e
was
a
signi ican
associa ion
be ween
he
occu ence
o
igh ly
olded
pinnae
and
o pidi y,
bu
ha
his
associa ion
was
s a is ical
and
no
absolu e,
as
was
emphasized
by
he
poo
ela ionship
be ween
e ec ion
o
he
pinnae
and
body
em-
pe a u e.
The e
was
also
no
ela ionship
be ween
he
ele a ion
o
body
empe a u e
abo e
ambien
and
ea
posi ion
o
ei he
o pid o
ac i e
ba s.
He
concluded
ha
pinna
posi ion
did
no
e lec
he mo egula o y
beha iou
and
sugges ed
ha ,
al hough
long-ea ed
ba s
may
old
hei
pinnae
o
conse e
hea
in
he
i s
place,
hey
may
ail
o
keep
hem olded
i
hey
begin
o
mo e
hei
wings
and
so
allow
he
ea s
o
un old.
Thus,
ac i i ies
such
as
g ooming
o
adjus ing
hanging
posi ion
may
be
incompa ible
wi h
keeping
he
pinnae
olded.
Since
o pid
ba s
seldom
mo e,
hey
a e
mo e
likely
o
keep
hei
ea s
olded,
and
hus
ea
posi ion
depends
p incipally
on
wha
he
ba
is
doing
and
no
on
how
wa m
o
cold
i
is.
Besides
educing
hea
loss,
a
second
possible
unc ion
o
pinna
olding
was
p oposed
by
Coles
e al.
(1989).
Because
he
d as ic
al e a ion
o
shape
in ol ed
in
olding
he
pinna
sig-
ni ican ly
educes
i s
acous ical
e iciency,
hey
sugges ed
ha
olding
may
be
used
o
educe
audi o y
sensi i i y
when
he
ba
is
inac i e
and
so
a oid
acous ical
dis u bance.
An
analogy
is
he
pa ial
closing
o
he
acial
u
in
sleeping
owls.
Speakman
e al.
(1991)
pu sued
his
a gumen
and
sugges ed
ha ,
by
olding
hei
ea s
du ing
o po ,
long-ea ed
ba s
educe
hei
sensi i i y
o
sounds
and
so
p e en
equen
and
unnecessa y
a ousals
which
would
was e
ene gy.
The
ailu e
o
non- ac ile
s imuli
o
p ecipi a e
a ousals
in
hibe na ing
ba s
(see
Chap e
8)
is
consis en
wi h
his
heo y.
Bekke
and
Mos e
(1990)
sugges ed
ha
long-ea ed
ba s
old
hei
ea s
in
o de
o
educe
he
likelihood
o
being
pulled
down
om
hibe na ion
si es
by
p eda o s,
and
ano he
sugges ion
(J.
Speakman,
pe s.
comm.)
is
ha
hey
old
hem
in o de
o
a oid
os bi e.
Func ions
o
La ge
Ea s
G i in
(1958)
compa ed
he
abili y
o
a
pleco ine
ba ,
Co yno hinus
ownsendii,
wi h
ha
o
Myo is
luci ugus,
a
ba
wi h
no mal
leng h
ea s,
o
a oid
wi es
a anged
in
a
g id
in
a
ligh
oom.
He
ound
ha
any
loss
o
e iciency
in
C.
ownsendii
caused
by
he
ain ness
o
i s
ul a-
sonic
calls
was
compensa ed
o
by
i s
abili y
o
hea
much
ain e
echoes
han
did
M.
luci ugus.

24
Recogni ion
and
Design
Fea u es
Howe e ,
i s
abili y
o
a oid
he
wi es
was
educed
i
he
shape
o
posi ion
o
he
pinna
was
al e ed
in
any
way.
The
la ge
pinnae
we e
hus
assumed
o
be
an
impo an
componen
in
he
abili y
o
la ge
ea ed
ba s
o
echoloca e
using
low-in ensi y
ul asound.
Coles
e
al.
(1989)
eco ded
he
neu al
audiog am
o
P.
au i us
and
ound
ha
he
mos
sensi i e
bes
equency
h esholds
o
single
neu ons
we e
be ween
7
and
20
kHz,
and
ha
he
lowe
and
uppe
hea -
ing
limi s
we e
3
kHz
and
63
kHz
espec i ely.
The
audi o y
sys em
o
his
species
is
hus
excep ionally
sensi i e
o
low- equency
sounds,
well
below
hose
used
in
echoloca ion.
The
pinna
was
ound
o
beha e
as
an
e icien
p essu e
ans o me
abo e
7
kHz,
due
o
i s
size
and
ho n-like
p ope ies,
and
he
ex e nal
ea
as
a
whole
o
p o ide
abou
20
dB
in
acous ic
gain
in
he
equency
ange
o
8-20
kHz
(Coles
e ai,
1989).
Such
ampli ica ion
o
sound
p essu e
is
likely
o
play
a
majo
ole
in
enhancing
low
equency
sensi i i y,
he
equency
ange
a
which
he
audi o y
sys em
o
P.
au i us
is
mos
sensi i e.
The
main
unc ion
o
he
huge
ea s
hus
appea s
o
be
connec ed,
no
wi h
echoloca ion,
bu
wi h
de ec ing
low- equency,
p ey-
gene a ed
sounds
such
as
mo h
lu e ing.
I
is
an
impo an
componen
in
he
echnique
o
gleaning,
which
is
discussed
in
Chap e
3.
Ob is
e
al.
(1993),
in
a
ield
s udy
in ol ing
47
ba
species
om
13
amilies,
simila ly
ound
ha
he
la ge
pinnae
o
ba s
which
used
p ey-
gene a ed
sounds
o
ind
hei
a ge s
supplied
high
sound
p essu e
gain
a
lowe
equencies.
D ag
E ec
o
La ge
Ea s
La ge
ea s
p oduce
a
high
deg ee
o
d ag
(No be g,
1976a),
hus
inc easing
he
amoun
o
ene gy
long-ea ed
ba s
ha e
o
expend
in
ligh .
Rayne
(1987)
ound
ha
d ag,
and
he e-
o e
h us
o
o e come
i ,
a ied
wi h
ligh
speed
-
he
as e
a
ba
lew,
he
g ea e
was
he
d ag
caused
by
i s
ea s.
No be g
(1976a)
obse ed
ha
P.
au i us
o en
lies
slowly
and
spec-
ula ed
ha
only
ba s
which
specialize
in
low-speed
ligh
can
a o d
o
ha e
la ge,
d ag-
p oducing
ea s.
Al hough
his
gene aliza ion
holds
ue
o
mos
species,
he e
a e
excep ions,
such
as
he
as - lying,
big-ea ed
molossid
O omops
(Hill
and
Smi h,
1984).
EYES
The
eyes
o
long-ea ed
ba s
a e
ela i ely la ge
compa ed
wi h
hose
o
all
o he
Eu opean
species.
C anb ook
(1963b)
specula ed
ha
sigh ,
as
well
as
hea ing,
may
be
used
o
loca e
p ey
while
gleaning.
I
is
emp ing
o
assume
ha
he
possession
o
big
eyes
means
Pleco us
species
see
be e
han
o he
ba s,
bu
in
ac
e y
li le
in es iga ion
has
been
ca ied
ou .
Eisen au
(1930)
conduc ed
isual
acui y
expe imen s
on
P.
au i us
and
se o ine
(Ep esicus
se o inus)
ba s
in
Ge many.
He
ained
he
ba s
wi h
ood
ewa ds
o
selec
ca ds
colou ed
black
o
whi e.
Bo h
species
we e
able
o
selec
9
cm
squa e
ca ds
o
he
app op ia e
colou
om
a
dis ance
o
5
cm,
bu
bo h
ailed
when
o e ed
a
choice
be ween
a
ci cle
and
a
c oss
o
simila
size.
I
does
no
appea
om
hese
expe imen s
ha
P.
au i us
has
pa icula ly
good
eyesigh .
Howe e ,
as
Ransome
(1990)
poin ed
ou ,
he
expe imen s
we e
ca ied
ou
in
b igh
ligh ,
condi ions
in
which
all
ba s
a e
likely
o
become
con used
and
which
hey
will
no
no mally
encoun e
in
he
wild.
Ransome
conside ed
ha
he
opic
equi ed
u -
he
in es iga ion
and
ad oca ed
expe imen s
unde
low
ligh
condi ions
and
using
a
ange
o
species.
Such
in es iga ions
may
disco e
whe he
Pleco us
ba s
can,
in
ac ,
see
be e
unde
hese
condi ions
han
can
ba s
wi h
smalle
eyes,
such
as
Rhinobphus
species.
Ch omosomes
25
NOSTRILS
Long-ea ed
ba s
a e
unusual,
bu
no
unique,
in
ha
hey
p oduce
ul asonic
calls
h ough
he
nos ils
and
no
h ough
he
mou h.
They
do
no
ha e
specialized
appendages
such
as
he
ho seshoe-shaped
noselea
o
he Rhinolophidae
o
p oducing
he
calls,
bu
hei
nos ils
a e
shaped
di e en ly
om
hose
o
mos
o he
Eu opean
es-
pe ilionid
species.
They
open
la e ally
and
appea
o
be
sli -shaped
a he
han
ound
as
a e
hose
o ,
o
example,
Myo is
species.
Howa d
(1995)
examined
he
nos ils
o
P.
au i us
in
de ail
and
eco ded
ha ,
al hough
hey
a e
appa en ly
elonga e,
only
he
an e-
io
pa
o
hem
opens
in o
he
nasal
passage.
The
pos e io
pa
opens
in o
a
ci cula
pi -like
s uc u e
sepa a ed
om
he
an e io
pa
by
a
sep um.
He
specula ed
ha
his
s uc u e
may
be
conce ned
wi h
he
p oduc ion
o
esonance
o
wi h
smell.
The
exac
way
in
which
ul asound
is
p oduced
by
he
nos ils
o
Pbco us
species
has
no
been
de e mined,
bu
i
seems
mo e
han
likely
ha
he
unusual
s uc u e
o
he
nose
is
con-
ce ned
wi h
his
unc ion.
Because
ul asound
is
no
p oduced
h ough
he
mou h,
long-
ea ed
ba s
a e
also
unusual
in
ha
hey
ly
wi h
hei
mou hs
closed.
When
ale
in
he
oos
and
p oducing ul asound
(e.g.
jus
p io
o
aking
ligh )
o
when
being
handled,
hey
also
keep
hei
mou hs
shu .
This
may
well
be
one
o
he
easons
why
hey
ha e
such
an
appealing
image
and
a e
seen
by
he
public
as
' iendly'
ba s
-
a
mou h ul
o
ee h
appea s
h ea ening
e en
i
i
is
caused
by
ba s
simply
'looking
a ound'
o
ind
ou
wha
is
going
on.
CHROMOSOMES
The
ka yo ypes
o
P.
au i us,
P.
aus iacus
and
B.
ba bas ellus
a e
iden ical
-
2N=
32
(Fedyk
and
Fedyk,
1970).
The e
a e
en
pai s
o
me acen ic
au osomes,
i e
pai s
o
elocen ic
au osomes,
a
subme acen ic
X
ch omosome
and
a
small,
ac ocen ic
Y
ch omosome
(Fedyk
and
Rup ech ,
1983).
FNa
=
50-54
in
P.
au i us
and
50-52
in
P.
aus iacus
(Zima
and
Ho acek,
1985).
Figu e
2.7
shows
a
ka yog am
o
a
male
P.
aus iacus
om
he
Czech
Republic.
1
2
3
4
5
6 7
8
9
10
11
12
13
14
15
X
Y
I
1
10
nm
2-7
Ka yog am
o
a
mab
P.
aus iacus
om
he
Czech
Republic
( ep oduced
wi h
pe mission
om
Bake ,
1970).
26
Recogni ion
and
Design
Fea u es
WINGS
AND
FLIGHT
Pleco us
ba s a e
gene ally
conside ed
o
be
slow- lying
and
manoeu able,
able
o
ho e
and
o
ly
and
o age
in
clu e ed
en i onmen s.
Because
P.
au i us
adap s
ela i ely
easily
o
cap i i y
and,
wi h
i s
abili y
o
ly
slowly,
is
sui able
o
expe imen s
in
ligh
unnels
and
chambe s,
i s
ligh
cha ac e is ics
ha e
been
s udied mo e
closely
han
hose
o
any
o he
Eu opean
species.
Da a
on
P.
aus iacus
a e
spa se,
bu
he
mo phology
and
beha iou
o
he
wo
species
a e
so
simila
ha
i
is
unlikely
ha
hey
di e
g ea ly
in
espec
o
how
hey
ly.
Wing
Mo phology
and
Shape
Figu e
2.8
shows
he
wing
s uc u e
o
a
Pleco us
ba .
The
p opa agium
is
he
sec ion
o
wing
which
lies
an e io
o
he hume us
and
adius
(Al ingham,
1996).
Wi h
he
dac y-
lopa agium
minus
(be ween
digi s
2
and
3),
i
ac s
as
a
wing
lap
on
he
leading
edge.
I
can
be
aised
o
lowe ed
using
he
humb
and
digi
2,
and
is
adjus ed
by
he
ba
o
p e en
s alling
a
low
speeds.
The
p opa agium
is
g ea es
in
a ea
in
slow- lying,
manoeu able
ba s
FIG
2.8
S uc u e
o
he
wing
o
Pleco us,
showing
he
pa s
o
he
pa agium.
Wings
and
Fligh
27
and
is
la ge
in
Pleco us
han
in
mos
Eu opean
espe ilionid
species.
Long-ea ed
ba s
also
ha e
ela i ely
b oad
wings
(measu ed
as
he
leng h
om
he
w is
o
he
end
o
digi
5).
The
a ea
o
wing
be ween
he
body
and
digi
5
( he
plagiopa agium)
p o ides
mos
o
he
li
in
ligh ,
and
hus
Pleco us
ba s
a e
able
o
gene a e
mo e
li
han
a e
many
o he
species.
Wing
a ea
is
impo an
when
dealing wi h
ligh
mo phology,
and
i
is
usually
mea-
su ed
by
d awing
a ound
he
en i e
ligh
memb ane
on
g aph
pape
and
hen
coun ing
he
squa es.
Wing
loading
is
de ined
as
weigh
o
ba
di ided
by
o al
a ea
o
ligh
memb ane
and
is
hus
a
measu e
o
he
load
ca ied
pe
uni
o
memb ane.
Aspec
a io
is
calcula ed
as
wingspan
di ided
by
a ea
o
ligh
memb ane.
I
can
be
used
o
compa e
he
leng h
o
wings
in
di e en
ba s
in
ela ion
o
wing
a ea.
As
a
ule,
sho ,
b oad
wings
(low
aspec
a io)
a e
associa ed
wi h
slow
ligh
(No be g,
1976a)
and
long,
na ow
ones
(high
aspec
a io)
wi h
as
ligh .
Wing
loading
a ec s
he
speed
a
which
ba s
ly
—
he
highe
he
wing
loading,
he
as e
he
ba
has
o
ly
in o de
o
gene a e
enough
li
o
s ay
ai bo ne.
Low
wing
loading
ba s
can
ly
mo e
slowly.
P.
au i us
has
an
a e age
wing
loading
alue
o
7.2
N
m~2
(0.072
g
cm"2)
and
an
aspec
a io
o
5.9
(No be g,
1970a;
1976b;
No be g
and
Rayne ,
1987).
In
P.
aus iacus,
wing
loading
is
7.9
N
n:
and
aspec
a io
6.1
(No be g
and
Rayne ,
1987).
Simila
alues
ha e
been
eco ded
o
o he
pleco ine
ba s:
wing
loading
has
been
calcula ed
a
7.0
N
m"2
in
C.
ownsendii
(Fa ney
and
Fleha y,
1969)
and
5.9
N
m
2
in
C.
a inesquii
(No be g
and
Rayne ,
1987)
and
aspec
a io
a
5.9
in
bo h
hese
species
(Fa ney
and
Fleha y,
1969;
No be g
and
Rayne ,
1987).
Long-ea ed
ba s
hus
ha e
low
wing
loading
alues
and
a e -
age
o
low
aspec
a ios
compa ed
wi h
o he
ba s
(No be g
and
Rayne ,
1987).
They
ha e
a
la ge
wing
a ea
and
b oad,
ela i ely
sho
wings.
This
combina ion
is
commonly
ound
in
ba s
which
eed
among
ege a ion,
since
i
allows
hem
o
make
igh
u ns
and
o
ho e .
Low
wing
loading
also
enables
such
ba s
o
ca y
hea y
p ey
i ems
o
pe ches
o
consume
hem.
Ano he
ea u e
ol
slow- lying,
igh - u ning
ba s
is
ha
hey
o en
ha e
sho ,
ounded
wing ips,
and
his
is
ue
o
bo h
Pleco us
species.
Also,
he
ips
o
digi s
3,
4
and
5
a e
ca -
ilaginous
and
lexible
(No be g,
1970b),
so
allowing
lexibili y
o
shape.
Finally,
long-
ea ed
ba s
ha e
high-cambe
wings
and
he
abili y
o
con ol
cambe
by
lexing
digi
5
and
lowe ing
he
hind
limbs.
No be g
and
Rayne
(1987)
conside ed
ha
he
e m
manoeu abili y
was
oo
o en
loosely
used,
and
hey
de ined
i
p ecisely
o
dis inguish
i
om
agili y.
Manoeu abili y
e e s
o
he
space
equi ed
o
a
ba
o
al e
i s
ligh
pa h
when
i
is
lying
a
a
cons an
speed.
I
is
g ea es
when
wing
loading
is
low
and
is
also
a ou ed
by
sho
wingspan;
Pleco us
ba s a e
hus
mo phologically
equipped
o
be
highly
manoeu able,
and
hei
abil-
i y
o
ly
and
u n
in
con ined
spaces
con i ms
ha
his
is
he
case.
Agili y
is
a
measu e
o
he
ease
o
speed
wi h
which
he
ligh
pa h
can
be
al e ed.
I
is
a ou ed
by
low
wing
ine -
ia
and
high
li
p oduc ion.
Some
as - lying
ba s
(e.g.
he noc ule,
Nyc alus
noc ula)
a e
agile
bu
no
manoeu able,
while
all
manoeu able
ba s
a e
also
agile.
S aigh
Fligh
and
Fligh
Speed
In
le el
ligh ,
he
long
axis
o
he
body
is
kep
almos
ho izon al
(No be g,
1976a).
Wing
and
body mo emen s
o
a
specimen
o
P.
au i us
lying
in
a
ne
cage
0.6
X
0.6
x
3.5
m
we e
28
Recogni ion
and
Design
Fea u es
ilmed
by
No be g
(1976a).
A
he
beginning o
he
downs oke,
he
wings
we e
ex ended
a
an
angle
o
49.4°
abo e
he
ho izon al
and
he
ail
and
ee
we e
held
s aigh
backwa ds.
By
he
middle
o
he
downs oke,
he
wing
was
wis ed
and
he
ee
and
ail
began
o
mo e
downwa ds,
and
in
he
la e
pa
o
he
downs oke
he
wings
we e
sha ply
cambe ed
and
he
ail
ully
lowe ed.
A
he
beginning
o
he
ups oke,
he
elbows
and
w is s
we e
sligh ly
lexed
and
he
cambe
s ill
p onounced,
bu
in
he
la e
pa
o
he
ups oke
he
ee
we e
aised,
which
educed
he
cambe
in
he
p oximal
pa
o
he
wings.
The
middle
pa
o
he
wings
was
s ill
cambe ed
and
he
wing ips
momen a ily
in e ed
be o e
beginning
he
nex
downs oke.
In
he
abo e
s udy,
he
ba
was
ound
o
be
lying
slowly,
a
an
es ima ed
speed
o
2.35
m
s~'.
No be g
calcula ed
ha , wi h
he
ba 's
huge,
d ag-p oducing
ea s,
he
speed
a
which
minimum
powe
was
equi ed
(i.e.
he
mos
ene ge ically
e icien
speed)
was
sligh ly
as e
han
his
a
3.1
m
s"1.
This
is
slow
o
a
ba ,
bu
co esponds
o
he
speeds
a
which
P.
au i us
has
equen ly
been
obse ed
lying
while
o aging
a ound
ege a ion.
No be g
(1976a)
sugges ed
ha
only
ba s
which
o age
a
slow
speeds
can
a o d
o
ha e
la ge
ea s,
since
he
coe icien
o
d ag
inc eases
apidly
as ligh
speed
inc eases.
Howe e ,
Pleco us
ba s
do
no
always
ly
so
slowly.
Baagoe
(1987)
used
ba
de ec o s,
nigh
ision
equipmen
and
mul iple- lash
pho og aphy
in ol ing
wo mo o -d i en
came as
a
ixed
dis ance
apa
o
measu e
ligh
speeds
o
12
Scandina ian
ba
species.
In
s aigh
o
wide- u ning
ligh ,
he
es ima ed
ha
P.
au i us
lew
a
an
a e age
speed
o
4.5
m
s"1.
This
was
almos
iden ical
o
he
ligh
speed
(4.6
m
s~')
o
Myo is
dauben onii,
a
ba
o
simila
size
and
wing
loading
o
7.0
N
m~2.
Nyc alus
noc ula,
a
la ge
ba
wi h
highe
wing
loading,
lew
as e
a
8.0
m
s~".
P.
au i us
had
he
slowes
speed
o
he
12
species,
bu
no
by
much.
Howa d
(1995)
simila ly
ound
ha
P.
au i us
commu ing
be ween
wo
ixed
poin s
along
a
ou e
be ween
oos
and
o aging
a eas
lew
a
5.9
m
s~
Again,
his
speed
is
compa able
wi h
commu ing
speeds
o
simila
sized
species
such
as
Myo is
na e e i
and
M.
dauben onii.
Thus
i
appea s
ha
P.
au i us
can
ly
ela i ely
as
and
ha
i
does
so
in
s aigh
ligh
when
commu ing.
I
p obably
does
his
in
o de
o
educe
he
isk
o
being
caugh
by
p eda o s,
bu
pays
a
p ice
because
such
apid
ligh
is
mo e
ene ge ically
expensi e
han
slowe
ligh .
This
could
be
one
o
he
easons
why
P.
au i us
o age
close
o
hei
oos s
and
why
mos
oos s a e
wi hin
0.5
km
o
decidu-
ous
woodland
(En wis le
e
al.,
1997).
I s
minimum
powe
speed
is
he
slowe
one
used
du ing
o aging.
Tu ning
Fligh
Rayne
and
Ald idge
(1985)
used
a
mic ocompu e
o
econs uc
a
h ee-dimensional
objec
om
images
aken
by
s ill
o
cine
came as
and
we e
able
o
illus a e
a
slow,
powe ed
u n
by
P.
au i us
in
a
ligh
chambe .
The
ba
gained
heigh
as
i
slowed
on
app oaching
he
u n
and
hen
decele a ed
du ing
he
u n
om
a
s aigh
ligh
speed
o
1.5
m
s"'
o
a
minimum
o
0.55
m
s"1,
when
he
cu a u e
was
s eepes .
The
wings
gene a ed
li
h ough-
ou
he
u n,
bu
did
no
p oduce
enough
e ical
o ce
o
suppo
he
ba 's
weigh
because
he
wings
had
o
be
banked
o
p oduce
he
la e al
cen ipe al
accele a ion
equi ed
o
al e
he
di ec ion
o
he
ligh
pa h.
In
ano he
s udy,
Ald idge
(1987)
in es iga ed
u ning
ligh
in
six
B i ish
ba
species
including
P.
au i us
using
Pennycuick's
(1975)
equa ion:
Wings
and
Fligh
29
217
=
;
p
CL
Sg
sin
0
whe e
p
is
he
ai
densi y,
CL
is
he
li
coe icien ,
S
is
he
wing
a ea,
g
is
he
accele a ion
due
o
g a i y,
W
is
he
weigh
o
he
ba
and
0
is
he
bank
angle.
F om
his
equa ion,
Ald idge
deduced
ha
he
u ning
adius
was
p opo ional
o
wing
loading
and
ha
ba s
wi h
low
wing
loading would
u n
igh ly
and
be
manoeu able.
The
ba s
we e
ained
o
ly
in
a
ligh
unnel
and hen
made
o
u n
by
a
ba ie
being
placed
ac oss
hei
ligh
pa h.
The
u ns
made
we e
pho og aphed
and
cu a u e,
speed
and
la e al
and
e ical
accele a-
ion
calcula ed.
The
li
coe icien
o
each
species
was
es ima ed
om
he
equa ion
abo e;
hese
we e
he
coe icien s
which
would
ha e
de eloped
i
he
ba s
had
been
pe o ming
gliding,
a he
han
powe ed,
u ns.
High
alues
indica ed
ha
powe ed
u ns
mus
ha e
been
used,
and
a
Cj
alue
o
1—2
was
equi ed
o
indica e
ha
a
ba
was
gliding.
P.
au i us
was
he
only
species
wi h
a
CL
in
his
ange
(CL
=
1.57).
Thus
i
was
essen ially
pe o ming
an
unpowe ed,
o
gliding,
u n.
The
o he
ba s,
wo
Rhinolophus
species,
wo
Myo is
species
and
P.
pipis ellus,
all
pe o med
powe ed
u ns
and
app oached
he
u n
a
ela i ely
high
speeds.
P.
au i us
can
he e o e make
igh ,
powe ed
u ns
o
gliding
u ns
and
is
able
o
do
bo h
a
ela i ely
low
ene ge ic
cos
because
o
i s
low
wing
loading.
Rhinolophus
e ume-
quinum
was
also
ound,
in
he
s udy,
o
be
able
o
make
igh u ns,
bu
a
highe
ene gy
cos .
Low
ene gy
cos
is
hus
a
ea u e
o
wis ing,
u ning
ligh
in
P.
au i us.
Ho e ing
in
Pleco us
species
has
been
in es iga ed
in
a
numbe
o
s udies
since
i
was
i s
desc ibed
by
Eisen au
(1936).
No be g
(1970a)
ilmed
a
ho e ing
P.
au i us
in
a
ligh
unnel
and
measu ed
wingbea
equency
a
10.2—12.5
Hz.
The
mean
speed
o
he
wing ip
was
abou
5.2
m
s~'
a e aged
o e
an
en i e
cycle.
The
g ea es
speed
o
he
wing ip
was
eached
in
he
uppe
hal
o
he
ups oke and
in
he
middle
o
he
downs oke.
The
wings
swep
downwa ds
and
o wa ds
ully
ex ended
and
wi h
high
cambe
and hen
u ned
upwa ds
and
sligh ly
owa ds
each
o he a
he
bo om
o
he
s oke.
The
ups oke
s a ed
wi h
a
sligh
lex
o
he
elbow
and
w is
and
o a ion
o
he
hume us,
which
inc eased
he
angle
o
a ack.
The
o elimb
ose
upwa ds
and
backwa ds
un il
he
adius
was
inclined
a
60-70
o
he
ho izon al,
a
which
poin
a
simul aneous
e e sing
o a ion
o
he
hume us
caused
he
wing
o
lick
backwa ds
and
upwa ds
be o e
he
s a
o
he
nex
downs oke.
Du ing
he
cycle
he
u opa agium
mo ed
up and
down
as
a
esul
o
leg
mo emen s;
his
had
he
e ec
o
keeping
a
ai ly
cons an
angle
o
a ack
on
he
p oximal
pa
o
he
wing
and
also
main ained
he
equilib ium
o
he
body
du ing
he
cycle.
The
long
axis
o
he
body
was
inclined
a
abou
30°
o
he
ho izon al,
and
he
body
lexed
and
ex ended
sligh ly
du ing
he
cycle;
i
was
mos
lexed
du ing
he
i s
hal
o
he
ups oke.
The
li
coe icien
ob ained
by
P.
au i us
du ing
ho e ing
is
3.1
o
6.4
(No be g,
1976b)
and
ho e ing
is
he e o e
mo e
powe -demanding
han
ei he
s aigh
o
u ning
ligh .
No be g
and
Rayne
(1987)
classed
P.
au i us
and
P.
aus iacus
as
ho e -gleaning
ba s,
i.e.
ba s
which
glean
om
su aces
such
as
ege a ion
o
ee
unks
and
which
ho e
b ie ly
beside
hese
su aces.
They
poin ed
ou
ha ,
o
ene ge ic
easons,
ho e ing
ba s
should
bene i
om
long
wings
bu
ha ,
because
ho e ing
akes
up
ela i ely
li le
o
long-ea ed
ba s
ime,
selec ion
has
ac ed
agains
he
e olu ion
o
long
wings
because
hese
would
30
Recogni ion
and
Design
Fea u es
hampe
non-ho e ing
ligh
in
clu e ed
si ua ions.
Pleco us
he e o e
comp omise
wi h
a e age-leng h
wings,
low
wing
loading
and
ounded
wing ips,
hus
achie ing
good
manoeu abili y
a
he
cos
o
ela i ely
expensi e
ligh .
ECHOLOCATION
All
mic ochi op e an
species
use
ul asound
o
na iga e
and
o
de ec
hei
p ey
and
many
also
use
ei he
audible
o
ul asound
(o
a
combina ion)
o
social
communica ion.
The
e-
quency
ange
o
na iga ion
calls
in
Eu opean
species
is
18-120
kHz
and
many
o
hem
call
a
a ound
40-60
kHz.
High- equency
sounds
ha e
sho
wa eleng hs,
and
he
bes
sound
o
de ec ing
an
objec
is
one
whose
wa eleng h
is
a ound
he
same
leng h
as
ha
objec .
A
sound
wi h
a
equency
o
50
kHz
has
a
wa eleng h
o
6.8
mm
(Al ingham,
1996),
which
is
an
a e age
leng h
o
an
insec .
This
explains
why
so
many
ba s
call
a
a ound
50
kHz.
Ba s
emi
ul asound
in
pulses,
and
i
is
he
cha ac e is ics
o
hese
pulses
which
allow
iden-
i ica ion
using
ba
de ec o s.
The
pulses
may
be
equency
modula ed
(FM)
o
cons an
equency
(CF),
o
hey
may
be
a
combina ion
o
he
wo.
FM
pulses
sweep
downwa ds
h ough
a
ange
o
equencies
(usually
om
a ound
65
o
25
kHz),
while
CF
calls
a e
a
one
equency,
o en
wi h
a
sho
FM
componen
a
he
beginning
o
end.
FM
pulses
a e
ypically
sho
-
0.5-5
ms
-
which
allows
he
ba
o
inish
emi ing
a
pulse
be o e
he
echo
o
ha
pulse
comes
back
o
i .
F om
he ime
aken
o
he
echo
o
e u n,
he
dis ance
o
a
p ey
i em
can
be
es ima ed.
As
he
ba
app oaches
an
insec ,
pulses
become
sho e ,
and
in
he
e minal
phase
o
an
a ack,
he
pulse
a e
inc eases
apidly
un il
each
pulse
may
be
only
a
ac ion
o
a
millisecond
long.
This
is
he
so-called
' eeding
buzz'
which
can
clea ly
be
hea d
on
ba
de ec o s,
al hough
i
sounds
simila
o
all
species
and
so
canno
be
used
o
iden i ica ion.
The
echoloca ion
calls
o
bo h
P.
au i us
and
P.
aus iacus
a e
cha ac e ized
by
hei
quie ness
combined
wi h
a
e y
as
pulse
a e
(Ca o,
1994).
The
call
o
P.
au i us
was
analysed
by
Ahlen
(1981),
who
desc ibed
i
as
a
ain
and
sho
FM
sweep,
abou
2
ms
long
and
wi h
p ominen
second
ha monics.
The e
we e
se e al
peaks
in
he
equency
spec-
um,
he
highes
a
26,
42
and
59
kHz.
The
epe i ion
a e
was
20
pulses
pe
second
o
mo e.
Figu e
2.9
shows
a
diag amma ic
ep esen a ion
o
his
cha ac e is ic
call.
The
call
can
mos
clea ly
be
hea d
wi h
he
de ec o
se
a
45-50
kHz,
bu
i
is
e y
ain
and
is
only
de ec able
i
he
ba
is
less
han
abou
5
m
away.
This
quie ness
(o
absence
o
a
call
i
he
ba
is
u he
away
bu
isible),
combined
wi h
he
as
epe i ion
a e
and
he
obse ed
slow
ligh
and
la ge
ea s,
is
used
o
iden i y
a
long-ea ed
ba
in
ligh .
P.
au i us
is
no
always
a
whispe ing
ba .
Ahlen
(1981)
also
iden i ied
a
loud
pulse,
de ec able
a
40
m
o
mo e,
7.1
ms
long
and
sliding
in
equency
om
42
kHz
o
12
kHz,
ending
wi h
a
sho
(1
ms)
CF
componen
a 12
kHz
and
hen
a
e y
sho
downwa d
sweep
(Figu e
2.10).
This
call
is
hea d
on
a
de ec o
as
a
so
bu
powe ul
smack,
and
he
epe i ion
a e
is
abou
5.4
pulses
pe
second.
I
may
be
emi ed
in e mi en ly,
usually
when
he
ba
is
lying
inside
s uc u es
such
as
ba ns
o
mines,
o
mo e
egula ly
when
ou
in
he
open
(Ahlen,
1981).
Coles
e
al.
(1989)
no ed
ha
his
loud
long-sweep
had
i s
peak
equency
a
a ound
12
kHz,
which
is
igh
in
he
middle
o
he
mos
sensi i e
hea ing
ange
o
P.
au i us
and
specula ed
ha
i
may
well
be
a
long-dis ance
communica ion
call,
al hough
i s
exac
unc ion
is
unknown.
Echoloca ion
31
120
100
pulses
s
1
-m
80
60
40
20
0
L
_J
L_
20
40
Time
(ms)
60
FIG
2.9
Diag amma ic
ep esen a ion
o
he
cha ac e is ic
call
o
P.
au i us
( ep o-
duced
wi h
pe mission
om
Ca o,
1994).
120
100
80
60
40
20
-
pulses
s"
20
40
Time
(ms)
60
FIG
2.10
Diag amma ic
ep esen a ion
o
he
loud-Long
sweep
(Ahlen,
1981)
o
P.
au i us.
The
no mal
ul asonic
calls
p oduced
by
long-ea ed
ba s
a e
adap ed
o
hei
o aging
me hods,
whe eby
hey
ly
in
clu e ed
si ua ions
and
glean
p ey
om
ege a ion.
In
gen-
e al,
CF
signals,
which
inc ease
he
ange
o
a ge
de ec ion,
a e
used
by
ba s
which
hun
in
open
si ua ions,
while
FM
signals
allow
ba s
accu a ely
o
loca e
and
iden i y
obs acles
32
Recogni ion
and
Design
Fea u es
and
a ge s
(Ro e ud,
1987)
and
a e
he e o e
sui able
o
use
in
clu e ed
en i onmen s.
Gleane s,
which
mus
dis inguish
be ween
echoes
om
po en ial
p ey
and
hose
om
back-
g ound
ege a ion,
use
low
in ensi y,
sho
(<1
ms),
mul iha monic
FM
signals
(Ro e ud,
1987).
Pleco us
ba s
do
no
ha e
o
de ec
p ey
a
any
g ea
dis ance,
and
by using
quie
calls
hey
also
a oid
ale ing
insec s
which
can
hea
hem
(see
Chap e
3)
o
as
long
as
possible.
The
ene ge ic
cos
o
p oducing
echoloca ion
calls
is
ela i ely
high
in
e es ial
mam-
mals
and
es ing
ba s,
bu
in
lying
ba s
he e
is
e ec i ely
no
ex a
cos
abo e
ha
o
lying
because
he
calls
a e
made
by
he
same
muscles
which
lap
he
wings
(Speakman
and
Racey,
1991;
Speakman,
1993).
This
was
sugges ed
by
Speakman
and
Racey
(1991)
o
be
he
ea-
son
why
echoloca ion
has
no
e ol ed
in
many
g oups
o
e es ial
mammals.
Jones
(1994)
calcula ed
ha ,
in
gene al,
insec i o ous
ba s
p oduce
abou
one
call
pe
wingbea
in
he
sea ch
phase
o
an
insec
chase,
and
hus
he
p oduc ion
o
calls
cos s
no
mo e
in
ene ge ic
e ms
han
would
lapping
wi hou
calling.
Howe e ,
abou
12%
o
species
(almos
all
gleane s)
ade
in ensi y
o
epe i ion
a e
and
use
each
wingbea
o
p oduce
se e al
e y
low-in ensi y
calls
ins ead
o
one
loud
one.
Ba s
such
as
Pleco us
species
ha e
hus
adap ed
he
me hod
o sui
hei
own
o aging
beha iou .
CHAPTER
3
Gleaning
and
Die
T
HE
ela ionship
be ween
ba s
and
hei
insec
p ey
is
a
complex
one
which
e ol es
con-
inuously.
The
p esen
chap e
e iews
he
esponse
o
insec s
o
p eda ion
by
ba s
and
i s
implica ions
o
he
way
in
which
long-ea ed
ba s
ca ch
hei
p ey.
The
hun ing
me h-
ods
o
he
ba s a e
conside ed,
as
is
he
e ec
hese
ha e
on
he
composi ion
o
he
die .
INSECT
RESPONSES
TO
ECHOLOCATION
Echoloca ing
ba s
i s
appea ed
in
he
Eocene,
by
which
ime
insec s
we e
long
es ablished
(Sales
and
Pye,
1974),
and
g oups
such
as
mo hs
(Lepidop e a)
and
lacewings
(Neu op e a)
hen
esponded
o
he
h ea
posed by
he
new
p eda o s
by
de eloping
o gans
which
could
de ec
he
ba s'
ul asound.
The
p esence
o
hea ing
o gans
in
insec s
has
been
known
o
some
ime
-
as
long
ago
as
1877,
Whi e
sugges ed
ha
mo hs
migh
be
able
o
a oid
ba s
by
lis ening
o
hem
squeaking.
He
was
e e ing
o
he
audible
c ies
o
he
ba s,
since
a
ha
ime
echoloca ion
was
no
unde s ood,
bu
he
idea
was
la e
subs an ia ed
by
he
dis-
co e y
ha
mo hs
possess
ea s,
o
ympanic
o gans,
which
a e
sensi i e
o
ul asound
in
he
equency
ange
p oduced
by
ba s.
The
ympanic
o gans
o
noc uid
mo hs
we e
e iewed
in
de ail
by
Roede
(1967)
and
since
hen
simila
o gans
ha e
been
desc ibed
in
i e
o he
lepidop e an
amilies,
as
well
as
in
g een
lacewings
(Mille ,
1970),
some
bee les
(Fo es
e
al.,
1997),
man ises,
cicadas
and
wa e
boa men
and
one
amily
o
Dip e a
(Fulla d
and
Yack,
1993).
Lepidop e a
The
supe amily
Noc uoidea
includes
mo hs
o
he
amilies
Noc uidae,
No odon idae
and
A c iidae,
all
o
which
ha e
simila
hea ing
o gans
si ua ed
on
he
hi d
ho acic
segmen ,
behind
and
below
he
a achmen
o
he
second
pai
o
wings
(Sales
and
Pye,
1974).
Each
o gan
lies
wi hin
a
deep,
scale- ee
ecess
on
he
pos e io
wall
o
he
hi d
ho acic
segmen ;
he
s uc u e
was
desc ibed
by
Roede
and
T ea
(1937).
A
anspa en
ympanic
mem-
b ane
(equi alen
o
an
ea d um)
aces
obliquely
backwa ds
and ou wa ds
in o
he
ecess.
An e io
o
he
ympanic
memb ane
is
an
ai
sac,
an
expanded
pa
o
he mo h's
espi a-
o y
sys em,
ac oss
which
senso y
cells
a e
suspended.
The e
a e
wo
acous ic,
o
A,
cells,
which
a e
modi ied
bipola
neu ones.
These
lie
close
oge he ,
suspended
in
a
s and
o
is-
sue
which
uns
om
he
cen e
o
he
ympanic
memb ane
owa ds
a
skele al
suppo
33

34
Gleaning
and
Die
p ojec ing
in o
he
ai
sac.
The
p oximal
ne e
ib es
om
each
A
cell
un
in
he
issue
s and
owa ds
his
suppo
and
a e
joined
by
a
hi d
ne e
ib e
om
a
la ge,
non-acous ic
B
cell
close
o
he
suppo .
The
h ee
ib es
join
o
o m
he
ympanic
ne e
which
joins
he
main
ne ous
sys em
a
he
p e o ho acic
ganglion.
The
A
cells
a e
sensi i e
o
ib a ion
om
he
ympanic
memb ane
and
hey
gene a e
ne e
impulses
which
a e
ansmi ed
along
he
ympanic
ne e
o
he
cen al
ne ous
sys em.
When
app oached
by
a
ba , noc uid
mo hs
will
show
a
a ie y
o
esponses
including
loops,
spi als
and
changes
o
speed
o
di ec ion.
Roede
(1962)
obse ed
402
encoun e s
be ween
ba s
and
noc uid
mo hs
and
ound
ha
when
he
mo hs
eac ed
in
his
way,
only
7%
o
hem
we e
caugh ,
compa ed
wi h
50%
o
hose
which
did
no
espond.
Roede
and
T ea
(1969)
simila ly
es ima ed
ha
insec s
which
can
hea
ha e
up
o
40%
less
chance
o
being
cap u ed
by
ba s
han
ha e
hose
which
canno .
A
se ies
o
expe imen s
by
Agee
(1967,
1969)
s udied
he
eac ion
o
noc uid
mo hs
o
pulsed
ul asound.
A
pulse
a es
o
mo e
han
15
s~'
and
a
sound
p essu e
le els
highe
han
80
dB,
he
mo hs
esponded
by
looping
o
di ing
o
he
g ound.
Some
esumed
ligh
a e
2-4
s,
bu
o he s
emained
s a-
iona y
on
he
g ound
o
up
o
10
min.
I
he
sound
p essu e
was
lowe
han
80
dB,
he
mo hs
made
a
di ec ional
u n
away
om
he
sou ce
o
he
sound.
A
pulse
a es
o
less
han
2
s
',
hey
showed
no
di ec ional
esponse,
bu
he
ligh
pa e n
became
e a ic.
The
ympanic
o gans
o
No odon idae
and
A c iidae
a e
sensi i e
o
sounds
o
16-20
kHz,
wi hin
he
human
hea ing
ange,
and
also
o
hose
well
in o
he
ul asonic
ange;
hey
will
espond
o
pulse
a es
o
up
o
45
s~'
(Haskell
and
Bel on,
1956).
Some
a c iid
species,
besides
being
able
o
hea
ba s,
a e
also
able
o
'jam'
hei
calls.
Dunning
and
Roede
(1965)
showed
ha
se e al
a c iid
mo h
species
p oduced
s eams
o
clicks
in
esponse
o
ba
calls.
The
dominan
equency
o
hese
was
abou
60
kHz,
and
i
eco dings
o
hem
we e
played
o
lying
ba s,
he
ba s
u ned
away
om
he
sound.
Achy a
and
Fen on
(1992)
p oposed
ano he
unc ion
o
hese
clicks.
Many
a c iid
mo hs
a e
unpala able
and
ha e
wa ning
colo a ion.
Achy a
and
Fen on
sugges ed
ha
he
clicks
a e
used
as
aposema ic
signals
o
wa n
he
ba s
o
he
mo h's
unpleasan
as e.
A
ba
which
has
once
hea d
he
clicks
and
hen
ied
o
ea
he
mo h
is
unlikely
o
epea
he
expe ience.
Such
sys ems
do
no
always
wo k,
howe e
-
Thompson
(1982)
ound
a
ga den
ige
mo h
(A c ia
caja),
an
unpala able,
wa ningly colou ed
a c iid,
unde
a
P.
au i us
eeding
pe ch.
I
had
been
killed
by
he
ba
and
hen
ejec ed
whole.
Ob iously
his
pa icula
ba
had
s ill
o
lea n
i s
lesson.
Mo hs
o
he
supe amilies
Py aloidea
and
Geome oidea
ha e
ympanic
o gans
on
he
abdomen,
usually
on
he
i s
segmen
(Fulla d
and
Yack,
1993),
whe eas
a
ew
species
o
opical
Sphingoidea
(hawk
mo hs)
ha e
hem
on
he
labial
palps.
Recen
wo k
on
species
o
geome id
mo hs
which
eme ge
in
au umn
and win e
(Rydell
e
al.,
1997)
showed
ha
males
had
good
hea ing
b oadly
uned
o
25-40
kHz,
despi e
being
p eyed
on
less
by
ba s
han
a e
mo hs
which
ly
du ing
summe .
They
esponded
o
pulsed
ul asound
by
al e ing
hei
ligh pa h
o
di ing
owa ds
he
g ound.
Howe e ,
he
emales
o
hese
species,
which
a e
ligh less
and
he e o e
no
subjec ed
o
ae ial
p eda ion,
we e
ound
o
ha e
g ea ly
educed
ympanic
o gans
and
o
be
i ually
dea .
Neu op e a
One
amily
wi hin
he
Neu op e a,
he
g een
lacewings
(Ch ysopidae),
ha e
a
swelling
nea
he
base
o
each
o ewing.
In
he
genus
Ch ysopa
hese
swellings
con ain
senso y
cells
Ba s'
Responses
o
Insec
Hea ing
O gans
35
esembling
he
ympanic
ea s
o
mo hs;
hey
a e
sensi i e
o
equencies
o
up
o
100
kHz
(Mille ,
1970).
Mille
(1971)
epo ed
ha
he
esponse
o
g een
lacewings
o
a ihci
ul asound
pulses
was
a
eac ion
o
he
lexo
muscles
o
he
o ewing.
This
caused
e
insec
o
old
i s
wings.
The
beha iou al
esponse
began
40
ms
a e
s imula ion
(Mi
e ,
1975),
which
enabled
he
lacewing
o
di e
s eeply
ou
o
he
pa h
o
a
hun ing
a .
Coleop e a
Hea ing
o gans
ha e
been
desc ibed
in
some
species
o
ige
bee les
(Cicindehdae;
Spangle ,
1988)
and
dung
bee les
(Sca abaeidae;
Fo es
e
aL
1997).
The
o gans
di e
in
s uc u e
and
placemen
be ween
he
wo
g oups
and
ha e
e ol ed
qui e
sepa a ely,
hose
in
sca abaeids
consis
o
a
hin
ympanic
memb ane
backed
by
an
ai
sac
and
a e
loca e
behind
he
head
(Fo es
e
aL,
1997),
while
hose
o
ige
bee les
a e
on
he
i s
abdomi-
nal
segmen
(Spangle ,
1988).
O he
G oups
O he
insec s
epo ed
o
possess
hea ing
o gans
include
some
species
o
man is
(Yage
and
Hoy,
1986),
ield
c icke s
(Pollack,
1994),
Hemip e a
such
as
some
wa e
boa men
(Co ixidae)
and
cicadas
(Cicadidae)
and
one
amily
o
cyclo haphan
lies
(Fulla d
and
Yack,
1993).
Insec
ea s
a e
hus
polyphyle ic,
ha ing
e ol ed
many
imes
in
a
num
e
o
di e en
g oups.
This
sugges s
ha
hey
a e
an
impo an
ea u e
o
noc u nal
insec s
an
ha
p eda ion
by
ba s
exe s
s ong
e olu iona y
p essu e
on
such
insec s.
The
ea s
o
ympanic
insec s
ha e
hei
maximum
sensi i i y
in
he
ul asonic
ange
used
by
mos
o
he
ba s
likely
o
p ey
on
hem
(Fen on
and
Fulla d,
1979).
An
in e es ing
example
occu s
in
Hawaii
(Fulla d,
1987),
whe e
Lasiu us
cine eus
semo us
is
he
only
ba
species.
The
ea s
o
mo hs
on
he
islands
we e
ound
by
Fulla d
o
be
b oadly
uned
o
20-50
kHz
bu
o
be
less
sensi i e
han
mo hs
elsewhe e
o
he
highe
end
o
his
ange,
c.
semo us
uses
o ien a ion
calls
o
27.8
kHz,
and
he
mo hs'
maximum
sensi i i y
was
a
a ound
30
kHz.
_
.
a
,
The
ange
a
which
a
hun ing
ba
can
be
de ec ed
by
an
insec
will
s ongly
a ec
he
chance
ha
insec
has
o
a oid
cap u e
and
will
a y
wi h
he
in ensi y
o
he
ba
s
call.
A
a
dis ance
o
10
cm
om
a
ba 's
ace,
he
in ensi y
o
calls
anges
om
mo e
han
110
dB
o
65
dB
(Fen on
and
Fulla d,
1981),
and
ba s
can
de ec
insec s
a
up
o
20
m.
Se e al
species
o
mo h
can,
howe e ,
de ec
ba s
a
up
o
40
m
(Roede ,
1967).
In
go ie
,
e e
o e,
i
is
likely
ha
a
mo h
should
de ec
a
hun ing
ba
be o e
i
is
'spo ed
by
ha
ba .
Howe e ,
ba s
use
a
numbe
o
s a egies
o
inc ease
hei
chances
o
ge ing
ahead.
BATS'
RESPONSES
TO
INSECT
HEARING
ORGANS
Ba s
can
inc ease
he
speed
a
which
hey
ly.
One
example
is
Eude ma
macula um,
which
is
also
an
excep ion
o
he
gene aliza ion
ha
ba s
wi h
big
ea s
a e
gleane s.
This
species
has
eno mous
ea s
and
eeds
almos
exclusi ely
on
mo hs
(Ross,
1967),
bu
Leona d
an
Fen on
(1983),
in
a
de ailed
s udy,
ound
no
e idence
ha
i
gleaned.
Ins ead
i
lew
a
a
heigh
o
abou
10
m
and
caugh
mo hs
in
high-speed
di es
ending
almos
a
g ound
le el.
36
Gleaning
and
Die
I
appea ed
o
be
ollowing
he
mo hs
as
hey
ook
e asi e
ac ion
ha ing
hea d
i
coming.
Possibly,
by
using
hese
di es,
E.
macula um
is
able
o
o e come
he
ene ge ic
expense
o
ha ing
o
ly
as
despi e
he
d ag
caused
by
i s
big
ea s.
A
second
possible
s a egy
by
ba s
is
o
use
highe
echoloca ion
call
equencies
(Fulla d,
1992).
An
A ican
species,
Cloeo us
pe ci ali,
which
uses
he
highes
known
equency
o
any
ba
(212
kHz;
Fen on
and
Bell,
1981),
has
been
shown
o
eed
exclusi ely
on
mo hs
(Whi ake
and
Black,
1976).
In
Eu ope,
g ea e
and
lesse
ho seshoe
ba s
Rhinolophus
e -
umequinum
and
Rhinolophus
hipposide os
also
ea
many
mo hs
(Beck,
1995;
Jones
e
al.,
1995)
and
use
highe
equencies
(cons an
equency
componen s
a
83
kHz
o
R.
e -
umequinum
and
113
kHz
o
R.
hipposide os;
Ca o,
1994)
han
do
espe ilionid
species.
Ba s
may
also
use
lowe
in ensi y
calls.
This
is
o en
associa ed
wi h
gleaning,
as
in
P.
au i us.
Low-in ensi y
ul asound
educes
he
ange
a
which
ba s
can
be
de ec ed
by
insec s
(Fulla d,
1992);
a
combina ion
o
low
in ensi y
and
high
equency
is
used
in
A ica
by
membe s
o
he
amilies
Nyc e idae
and
Megade ma idae
and
he e
is
e idence
ha
mo hs
a e
unable
o
de ec
he
p esence
o
hese
ba s
u he
away
han
abou
2
m
(Fen on
and
Fulla d,
1979).
In
addi ion,
because
mo hs
appea
o
be
unable
o
di e en ia e
be ween
dis-
an ,
loud
ba s
and
close-by,
quie
ones
(Fau e
e
al.,
1993),
hey
a e
less
likely
o
ake
quick
e asi e
ac ion
in
esponse
o
low
in ensi y
calls.
Howe e ,
some
ba s
which
use
low-in en-
si y
sound
and
which
glean
do
no
ea
mo hs
o
any
g ea
ex en
(e.g.
Myo is
na e e i
in
B i ain; Shiel
e
al.,
1991;
Swi ,
1997)
and
i
is
possible
ha
he
use
o
low
in ensi y
calls
is
p ima ily
a
s a egy,
no
o
educe
de ec ion
by
insec s,
bu
o
a oid
he
ba s
dea ening
hemsel es
when
lying
in
clu e ed en i onmen s
such
as
among
oliage.
The
ou h
way
in
which
ba s
can
educe
de ec ion
by
insec s
is
o
pick
p ey
o
oliage
and
o he
su aces,
i.e.
o
glean.
S a iona y
o
c awling
mo hs,
as
well
as
lying
ones,
a e
able
o
de ec
and
eac
o
ba
calls
(We ne ,
1981),
bu
hose
wi h
ho acic
ea s
will
be
less
sen-
si i e
in
such
si ua ions
because
hei
ympanic
o gans
a e
co e ed
by
hei
wings
and
so
he
sound
will
be
mu led.
Noc uid
mo hs
we e
ound
o
be
less
good
a
de ec ing
a
gleaning
ba ,
Myo is
sep en ionalis,
han
an
ae ial
hun e ,
Myo is
lucijugus,
and
we e
pa icula ly
poo
a
de ec ing
he
gleane
when
hei
ea s
we e
co e ed
(Fau e
e
al.,
1993).
E en
when
a
es -
ing
mo h
does
de ec
a
ba ,
i s
escape
possibili ies
a e
limi ed.
I
can
' eeze'
(We ne ,
1981)
bu
is
unlikely
o
ha e
ime
o
ake
ligh
and
di e
ou
o
he
way;
once
de ec ed
i
is
hus
less
likely
o
escape
han
is
a
lying
mo h.
F om
a ailable
e idence,
i
appea s
ha
mo hs
ha e
no
ye
e ol ed
a
e y
e icien
s a egy
o
a oiding
gleane s.
Fau e
e
al.
(1993)
sug-
ges ed
his
may
be
because
ela i ely
ew
ba
species
wo ldwide
(abou
10%)
a e
gleane s
and
hus
e olu iona y
p essu e
on
mo hs
o
coun e
he
s a egy
has
no
been
su icien ly
s ong.
They
iewed
gleane s
as
p eda o y
'chea e s'
in
ba -mo h
in e ac ions.
Howe e ,
his
a gumen
may
no
apply
in
Eu ope,
whe e
he e
a e
many
gleaning
ba s
(e.g.
Myo is
myo is,
M.
bly hii
and
M.
na e e i)
and
whe e
P.
au i us
is
one
o
he
commones
species.
GLEANING
Gleaning
is
de ined
as
cap u ing
p ey
c awling
o
a
es
on
solid
su aces
a he
han
in
ligh .
Ba s
may
glean
om
he
g ound
(e.g.
Myo is
myo is-,
A le az
and
Pe in,
1995;
A le az,
1996a)
o
g ass
(e.g.
Myo is
na e e i-,
A le az,
1996b),
o
may
swi ch
be ween
hese
subs a es
(A le az
and
Pe in,
1995)
depending
on p ey
a ailabili y.
They
may
glean
Gleaning
om
oliage,
mainly
ees
and
bushes,
as
does
P.
au i us,
al hough
his
species
is
also
known
o
ake
p ey
om
g ass
and
occasionally
om
he
g ound
(Shiel
e
al.,
1991).
Ano he
o m
o
gleaning
is
pe ch
hun ing,
which
is
o en
p ac ised
by
ba s
which
eed
on.small
mamm
s
o
e y
la ge
opical
insec s.
An
example
is
he
A ican
alse
ampi e,
Ca diode ma
co ,
which
ea s
la ge,
g ound-dwelling
bee les
and
cen ipedes
(Vaughan,
1976).
This
ba
scans
an
a ea
o
g ound
om
a
pe ch
and
makes
sho
(less
han
5
s)
ligh s
o
cap u e
p ey;
i
may
aligh
on
he
g ound
o
a
second
o
wo
be o e
e u ning
o
i s
pe ch.
Ad an ages
o
Gleaning
The e
a e
se e al
ad an ages
o
ba s
o
gleaning
a he
han
ae ial
cap u e.
One
o
hese,
as
has
been
discussed
abo e,
is
ha
i
is
a
s a egy
agains
which
mo hs
ha e
lew
de ences.
Mo hs
a e
ela i ely
la ge,
so -bodied
insec s
and
a e
he e o e
a
p o i able ood
sou ce,
in
gene al,
gleaning
ba s
ea
a
highe
p opo ion
o
mo hs
han
do
ae ial
hun e s.
Gleane s
a e
also
able
o
ly
la e
a
nigh
han
a e
ba s
such
as
Pipis elluspipis ellus
which
ely
o
mos
o
hei
ood
on
he
dawn
and
dusk
peaks
o
lying
insec s
(see
Chap e
4).
This
means
ha ,
pa icula ly
a
high
la i ude,
gleane s
do
no
ha e
o
isk
being
seen
and
cap u ed
by
p eda-
o s
on
ligh
e enings.
They
a e
less
dependen
on
ai
empe a u e
han
a e
ae ial
hun e s,
and
his
is
also
a
pa icula
ad an age
in empe a e
egions
whe e
ai
empe a u e
can
a y
om nigh
o
nigh .
Numbe s
o
lying
insec s
ha e
been
shown
o
be
signi ican ly
educed
on
nigh s
when
dusk
empe a u e
alls
below
10°C
(Rydell,
1989a).
By
eeding
on
non- ly-
ing
insec s,
gleane s
a e
able
o
o age
success ully
on
coole
nigh s.
A
ou h
ad an age
is
ha ,
by
gleaning,
ba s a e
able
o
inc ease
he
size
and
also
he
ange
o
insec s
which
hey
can
ca ch.
La ge
mo hs
can
be
handled
on
ege a ion
han
in
ligh
and
non- lying
p ey
such
as
spide s,
ca e pilla s
and
ha es men
can
be
included
in
he
die ;
hese
a e
ob iously
no
a ailable
o
ae ial
hun e s.
Ba clay
(1991)
s u
le
Myo is
species
o
simila
size
in
an
upland
a ea
in
Canada,
a
1350-2150
m
abo e
sea
le el
and
51°N,
whe e
nigh s
we e
cold
in
summe
and
he
dusk
insec
peak
sho .
He
ound
ha
M.
luci ugus,
an
ae ial
o age ,
did
no
p oduce
young
in
he
a ea,
since
no
nu se y
colonies
we e
ound
and
all
specimens
caugh
we e
males.
Females
hibe na ed
in
he
upland
a ea
bu
mo ed
o
lowe
al i ude
in
summe .
Howe e ,
M.
e o is,
a
gleane ,
did
o m
nu s-
e y
colonies
and
ea
young.
He
a ibu ed
he
di e ence
in
ep oduc i e
beha iou
o
he
o aging
me hods
o
he
wo
species
and
sugges ed
ha ,
because
M.
e o is
was
able
o
glean
some
o
i s
p ey,
i
was
able
o
mee
he
high
ene gy
demands
o
p oducing
and
ea ing
young
in
he
cool,
upland
a ea.
Gleaning
in
P.
au i us
Gleaning
as
a
o aging
echnique
by
his
species
has
been
es ablished
by
bo h
obse a ion
and
die a y
s udies.
Millais
(1904)
obse ed
a
long-ea ed
ba
aligh ing
on
esh
ca le
dung
and
ea ing
he
lies
(yellow
dung
lies,
Sca hophaga
s e co a ia)
which
in es ed
i .
Ba e -
Hamil on
(1910)
lis ed
a
numbe
o
obse a ions
o
oliage
gleaning,
and
Swi
and
Racey
(1983)
ma ked
indi idual
P.
au i us
wi h
e lec i e
ings
and
obse ed
hei
o aging beha -
iou
o e
a
numbe
o
nigh s.
The
ba s
we e
seen
o
ho e
in
ligh ,
o
u n
equen ly
and
o
disappea
among
he
ee
oliage
o
abou
20
s
be o e
eappea ing.
Die a y
s udies,
including
hose
by
Swi
and
Racey
(1983),
Rydell
(1989b)
and
Shiel
e
38
Gleaning
and
Die
al.
(1991),
con i m
he
p esence
in
he
die
o
a h opods
which
do
no
ly
and
mus
he e-
o e
ha e
been
caugh
by
gleaning.
These
include
spide s
(o de
A aneae),
ha es men
(o de
Opiliones)and
cen ipedes
(class
Chilopoda),
and
he
die
also
includes
diu nal
insec s,
such
as
b achyce an
lies,
which
a e
unlikely
o
ha e
been
caugh
in
ligh
a
nigh .
Bo h
Rydell
(1989b)
and
Shiel
e
al.
(1991)
es ima ed
ha
app oxima ely
40%
o
he
die
was
gleaned,
e en
i
i
we e
assumed
ha
all
mo hs
we e
caugh
in
ee
ligh .
In
ac ,
he
p opo ion
o
he
die
gleaned
is
almos
ce ainly
highe ,
as
many
mo hs
a e
also
caugh
in
his
way.
Ande son
and
Racey
(1991)
ound
ha ,
o
he
mo hs
aken
in
a
ligh
oom
by
cap i e
b own
long-ea ed ba s,
app oxima ely
hal
we e
caugh
by
gleaning
and
hal
by
ae ial
cap u e.
Ande son
and
Racey
(1991)
desc ibed
gleaning
in
hei
cap i e
P.
au i us
in
a
ligh
oom
measu ing
2.8
X
3.0
X
3.0
m,
in o
which
we e
eleased
li e
mo hs
o
species
known
o
be
included
in
hei
na u al
die
(Thompson,
1982).
Mo hs
we e
caugh
by
gleaning
in
36%
(#=53)
o
obse a ions
and
in
he
ai
in
44%
(n=41),
and
ba s
gene ally
began
hun ing
om
a
s a iona y
posi ion
a
a
pe ch.
Mo hs
we e
gleaned
mainly
om
e ical
su aces,
bu
also
om
he
ceiling
and
occasionally
om
he
loo .
Pu sui s
las ed
om
a
ew
seconds
o
abou
hal
a
minu e,
and
ligh
was
o en
wis ing.
The
ba s
in
he
abo e
expe imen s
we e
ollowed
as
closely
as
possible
by
a
mic ophone
a ached
o
a
cane
manipula ed
by
an
obse e .
The
mic ophone
was
connec ed
o
a
QMC
S200
ba
de ec o
in
b oadband
mode;
his
moni o ed
as
accu a ely
as
possible
echoloca-
ion
calls
p oduced
du ing
mo h
cap u es.
Ba s
cap u ing
mo hs
in
he
ai
p oduced
calls
in
89%
(»=23)
o
cases,
bu
du ing
gleaning
calls
we e
eco ded
in
only
29%
(»=8)
o
a acks.
Fu he mo e,
gleaning
a acks
we e
shown
o
be
signi icandy
mo e
success ul
(/><0.001)
when
calls
we e
absen ,
i.e.
when
echoloca ion
was
'swi ched
o
by
he
ba .
Ande son
and
Racey
(1991)
belie ed
i
was
unlikely
ha
isual
cues
we e
being
used
du -
ing
hese
non-echoloca ing
a acks,
since
ligh
le els
in
he
expe imen s
we e
e y
low.
I
was
also
ob ious
h oughou
he
expe imen s
ha
ba s
we e
only
a ac ed
o
mo ing
p ey
and
ha
s a iona y
mo hs
we e
igno ed,
e en
a
e y
sho
dis ances.
The
mo emen s
which
ale ed
ba s
we e
lu e ing,
walking
o
lying,
and
ba s
which
we e
ale ed
made
cha ac e is ic
lis ening
mo emen s
o
he
head
and
ea s.
A
second
se ies
o
expe imen s
wi h
cap i e
P.
au i us
(Ande son
and
Racey,
1993)
showed
ha
hese
ba s
we e
able
o
dis-
c imina e
be ween
lu e ing
and
non- lu e ing
mo hs;
hey
chose
lu e ing
ones
in
92.6%
o
ials
when
isual
and
ul asonic
cues
we e
a ailable
and
in
85.9%
o
ials
when
such
cues
we e
elimina ed.
The
ba s
we e
clea ly
able
o
disc imina e
on
he
basis
o
he
sounds
p oduced
by
he
lu e ing
mo hs.
Many
o
hese
sounds
ha e
a
equency
below
20
kHz,
and
his
is
he
equency
a
which
P.
au i us
hea ing
is
mos
sensi i e
(Coles
e
al.,
1989).
Ande son
and
Racey
(1993) he e o e
p oposed
ha
he
ba s
used
passi e
lis ening
o
p ey-
gene a ed
sounds
o
loca e
mo hs
du ing
mos
gleaning
a acks.
Al hough some
noc uid
mo hs
p oduce
ul asound
in
ligh as
a
esul
o
he
hindwings'
ma gins
ubbing
oge he
(Wa e s
and
Jones,
1994),
mos
o
hese
sounds
ha e
a
equency
o
a ound
46
kHz,
which
is
well
abo e
he
mos
sensi i e
hea ing
ange
o
P.
au i us.
Thus
hey
a e
no
likely
o
be
o
much
help
o
he
ba
in
loca ing
lying
mo hs
by
passi e
lis ening
(Wa e s
and
Jones,
1994)
and
he e o e
echoloca ion
is
used
in
mos
ae ial
a acks.
In
Ande son
and
Racey's
(1991,
1993)
expe imen s,
ba s
equen ly
ho e ed
in
on
o
lu e ing
mo hs
o
se e al
seconds
be o e
gleaning
hem.
While
ho e ing
in
on
o
po en-
ial
p ey,
hey
we e
mo e
likely
o
emain
silen
han
o
p oduce
echoloca ion
pulses
—
echoloca ion
calls
we e
eco ded
in
only
24%
o
sequences
in
which
ho e ing
occu ed
Die
39
(Ande son
and
Racey,
1993).
The
ho e ing
dis ance
(i.e.
he
dis ance
om
he
p ey
o
he
place
whe e
he
ba
chose
o
ho e )
was
ela i ely
cons an ;
i
did
no
di e
signi ican ly
be ween
sequences
wi h
and
wi hou
echoloca ion
and
i
may
ha e
been
ela ed
o
he
in en-
si y
o
p ey
sounds.
Howe e ,
he
mean
du a ion
o
ho e ing
was
signi ican ly
longe
(/>=0.04)
in
sequences
in
which
echoloca ion
was
no
used.
Ande son
and
Racey
concluded
ha
p ey-gene a ed
lu e ing
sounds
we e
su icien
o
ale
o aging
P.
au i us
o
he
p es-
ence
o
mo hs.
The
ba s
mainly
used
passi e
lis ening
o
hese
low- equency
(below
20
kHz)
sounds
o
loca e
p ey,
and
ho e ed
close
o
mo hs
while
lis ening
o
and
loca ing
hem.
Gleaning
by
P.
aus iacus
The
sho age
o
da a
on
o aging
in
g ey
long-ea ed
ba s
makes
i
di icul
o
es ablish
he
deg ee
o
which
hey
glean
hei
p ey.
They
ha e
been
seen
lying
e y
slowly,
ho e ing
and
u ning
equen ly
while
lying
among
oliage
and
inside
s ables
and
sheep olds
(Ba a aud,
1990).
This
au ho
epo ed
ha
ligh - agged
P.
aus iacus
cap u ed
insec s
inside
a m
buildings
and
he
also
obse ed
hem
lying
e y
close
o
ee
oliage.
A
single
adio- agged
indi idual
was
acked
by
Fluckige
and
Beck
(1995),
and
his
ba
lew
among
he
oliage
o
a
ches nu
ee;
i
was
assumed
o
be
gleaning
while
ou
o
sigh .
Die a y
s udies
indica e
ha ,
while
P.
aus iacus
undoub edly
does
glean,
i
does
so
o
a
lesse
ex en
han
P.
au i-
us.
Beck
(1995)
ound
emains
in
aecal
pelle s
o
cen ipedes
and
spide s,
as
well
as
hose
o
insec s
such
as
ea wigs
and
booklice
(Psocop e a)
which
a ely
ly,
bu
hese
we e
all
p e-
sen
in
e y
low
p opo ions.
The
ba s
a e
a
high
p opo ion
o
mo hs
and
bee les,
bu
i
is
unce ain
how
many
o
hem
we e
gleaned.
Baue o a
(1982)
conside ed
ha
he
die
o
P.
aus iacus
indica ed
i
hun ed
mainly
in
ee
ai
space.
A
single
labo a o y
s udy
(I.
Kaip ,
K.
Heblish
and
H.-U.
Schni zle ,
unpublished)
epo ed
ha
his
species
con inued
o
emi
echoloca ion
calls
while
aking
mo hs
om
lea es
and
hus
appea ed
no
o
use
passi e
lis-
ening.
I
may
use
a
di e en
gleaning
me hod
om
P.
au i us,
bu
he
whole
ques ion
o
gleaning
in
g ey
long-ea ed
ba s
equi es
u he
in es iga ion.
Ae ial
Cap u e
Ande son
and
Racey
(1991)
made
de ailed
obse a ions
on
he
cap u e
o
mo hs
by
P.
au i-
us
in
he
ai
as
well
as
by
gleaning.
In
he
majo i y
o
ae ial
cap u es
in
he
ligh
oom
(72%;
«=23),
he
mo h
was
seized
in
he
ba 's
mou h,
bu
he
u opa agium
(13%;
n=4)
and
wing
memb ane
(6%;
n=2)
we e
also
used.
The
u opa agium
was
used
as
a
scoop
o
ca ch
mo hs
which
we e
below
he
ba ,
and
he
wing
ips
we e
used
o
d aw
p ey
in
la e ally
be o e
ans-
e ing
i
o
he
mou h.
On
9%
o
occasions,
a
some saul
echnique
was used
by
he
ba .
Du ing
his
manoeu e,
he
ba
was
obse ed
o
oll
o wa ds
and
downwa ds
in
he
ai ,
d opping
b ie ly
ou
o
i s
ligh
pa h
as
i
en eloped
he
mo h
wi h
i s
body,
wings
and
ail
memb ane.
I
esumed
i s
ligh
once
he
mo h
had
been
ans e ed
o
he
mou h.
DIET
The
majo i y
o
insec i o ous
ba
species
ha e
been
shown
o
be
oppo unis ic
o age s
a he
han
specialis
p eda o s
on
pa icula
p ey
ca ego ies
(Kunz,
1974;
Fen on,
1995).
40
Gleaning
and
Die
Findley
(1993)
e iewed
much
o
he
li e a u e
on
he
s uc u e
o
ba
communi ies
and
concluded
ha
he e
was
li le
e idence
ha
such
communi ies
we e
o ganized
by
compe-
i ion
o
esou ces;
appa en
o e lap
among
species
in
a
habi a
was
likely
o
be
accoun ed
o
by
beha iou al
lexibili y
among
he
ba s.
A le az
and
Pe in
(1995)
ag eed
ha
mos
s udies
dealing
wi h
ophic
ecology
in
insec i o ous
ba s
ei he
showed
hem
o
be
oppo -
unis ic
o
else
ailed
o
in es iga e
ood
a ailabili y.
Thei
own
wo k
on
he
sibling
species
Myo is
myo is
and
M.
bly hii
in
Swi ze land
ound
ha
he
wo
species,
which
occupy
na -
ow
ophic
niches,
showed
no
ac i e
p ey
selec ion
wi hin
hose
niches.
Selec ion
has,
howe e ,
been
shown
by
g ea e
ho seshoe
ba s,
Rhinolophus
e umequinum
in
England
(Jones,
1990).
These
ba s
p eyed
selec i ely
on
mo hs
and
la ge
cha e s,
and
a oided
o he
insec s
when
mo hs
and
cha e s
we e
abundan ,
al hough
hey
ook
smalle ,
less
p o i able
p ey
a o he
imes.
The
sophis ica ed
echoloca ion
calls
o
ho seshoe
ba s
may
allow
hem
o
disc imina e
be ween
insec s
a
a
dis ance
and
so
o
selec
p o i able
p ey i ems;
espe -
ilionid
ba s,
which
use
a
di e en
echoloca ion
sys em,
may
no
be
able
o
do
his
so
accu-
a ely
and
his
may
be
one
eason
why
mos
eed
unselec i ely.
Fen on
(1995)
quo ed
Pleco us
species
as
an
example
o
he
lexibili y
shown
by
espe -
ilionid
ba s
du ing
o aging
—
hei
abili y
ei he
o
exploi
lying
insec s
o
o
glean
allows
hem
o
su i e
in
cool,
damp
a eas
whe e
insec
numbe s
may
be
e a ic.
Al hough
ea ly
die a y
s udies
based
on
analyses
o
insec
emains
unde
eeding
pe ches
indica ed
ha
P.
au i us
was
a
mo h
specialis
and
a e
li le
else,
mo e ecen
in es iga ions
ha e
shown
ha
he
species
is
lexible
in
i s
die
and
akes
a
a ie y
o
p ey.
Wha
i
ea s
depends
on
wha
is
a ailable,
which
in
u n
depends
on
habi a ,
wea he
condi ions,
ai
empe a u e,
ime
o
yea
and
e en
he
ime
o
nigh
a
which
o aging
akes
place.
I
ollows
ha
any
die a y
s udies
which
aim
o
es ablish
selec i i y
mus
also
conside
wha
p ey
is
a ailable
o
he
ba s.
Die
Analysis
Me hods
Remains
unde
eeding
pe ches
Long-ea ed
ba s
equen ly
ca y
hei
la ges
p ey
i ems
o
a
eeding
pe ch,
o
nigh
oos ,
whe e
hey
hang
up
in
o de
o
handle
and
consume
hem.
They
im
o
ha d
o
unpala -
able
pa s
o
insec s
such
as
wings
and
legs
and
hese
hen
accumula e
unde
he
pe ch,
om
whe e
hey
can
be
collec ed
and
analysed.
Because
he
as
majo i y
o
emains a e
o
la ge
mo hs,
analyses
in
he
pas
led
o
he
conclusion
ha
hese
ba s
a e
no hing
else,
since
smalle
p ey
i ems
which
we e
no
aken
o
he
pe ch
we e
ob iously
missed.
The
me hod
is
he e o e
o
limi ed
use
in
die
analysis,
bu
i
also
has
ad an ages.
Faecal
analysis
is
unable
o
dis inguish
be ween
di e en
mo h
species,
o
e en
amilies,
since
ew
mo h
emains
besides
scales
su i e
he
passage
h ough
he
ba 's
gu .
Because
emains
unde
pe ches
equen ly
include
whole,
o
almos
whole,
wings,
iden i ica ion
o
species
is
possi-
ble.
This
p o ides
aluable
in o ma ion
on
he
size
o
p ey
aken,
as
well
as
on
he
p obable
habi a
and
ime
a
which
hey
we e
caugh .
By
pai ing
wings,
in es iga o s
a e
also
able
o
assess
he
numbe
o
la ge
mo hs
which
ha e
been
consumed.
Faecal
analysis
This
is
he
mos
widely
used
me hod
o
in es iga ing
ba
die s.
Because
a h opods
ha e
an
exoskele on
made
o
chi in,
a
subs ance
only
pa ially
diges ible
by
animals,
pieces
o
Die
41
exoskele on
su i e
passage
h ough
he
gu
in
a
ecognizable
o m.
They
can
be
ex ac ed
om
he
aecal
pelle s
and
he
a h opods
hey
came
om
iden i ied,
usually
o
o de
and
o en
o
amily
o
e en
species.
Faecal
pelle s
a e
ela i ely
easy
o
collec ,
and
he
me hod
has
he
ad an age
o
no
ha ming
o
dis u bing
he
ba .
Resul s
a e
exp essed
as
ei he
pe -
cen age
occu ence
( he
pe cen age
o
pelle s
which
con ain
emains
o
a
p ey
ca ego y)
o
pe cen age
equency
( he
numbe
o
agmen s
o
a
ca ego y
-s-
he
o al
numbe
o
ecog-
nizable
agmen s
X
100).
The
me hod
is
ully
desc ibed
by
McAney
e al.
(1991).
The
only
o he
p ac ical
me hod,
analysis
o
s omach con en s,
in ol es
killing
he
ba
and
is
he e-
o e
unaccep able
since
ba
species
a e
legally
p o ec ed
in
Eu ope
and
all
ace
conse a ion
p oblems.
Faecal
analysis
is
ime-consuming
and
has
been
c i icized
as
being
inaccu a e
because
ba s
chew
hei
ood
ho oughly
and
end
o
im
o
he
pa s
o
insec s
which
a e mos
use ul
o
iden i ica ion.
Also,
ha d-bodied
insec s
such
as
bee les su i e
he
passage
h ough
he
gu
be e
han
do
so -bodied
ones
such
as
may lies
o
e y
small
ones
such
as
midges.
Howe e ,
he
abo e
wo
poin s
do
cancel
each
o he
o
some
ex en ,
since
ha d-
bodied
insec s
will
end
o
be
immed
mo e
by
ba s
han
so -bodied
ones
-
a
bee le
will
be
immed
mo e
han
a
midge
and
he e o e
midges'
legs,
an ennae
and
bi s
o
wing
a e
mo e
likely
o
be
ound
in
pelle s
han
a e
hose
o
bee les.
In
an
a emp
o
e alua e
he
eliabili y
o
aecal
analysis
in
de e mining
ood habi s,
Kunz
and
Whi ake
(1983)
ed
a ious
insec s
o
cap i e
ba s
and
hen
i s
aecal
pelle s
we e
analysed
by
a
esea che
who
had
no
p io
knowledge
o
he
die
composi ion.
The
ou
commones
axa
in
he
die
we e
iden i ied
in
he
co ec
o de
o
impo ance,
hus
demon-
s a ing
ha
he
me hod
was
alid
and
could
p oduce
eliable
esul s,
al hough
some
limi-
a ions
we e
e ealed.
Insec
Sampling
Me hods
Ligh
aps
Ligh
aps
a e
easy
o
anspo ,
inexpensi e
and
easy
o
use
in
he
ield.
They
consis
o
a
ligh ,
usually
me cu y
apou
o
ul a iole ,
abo e
a
collec ing
ub.
Thei
disad an age
is
ha
hey
ca ch
only
pho o ac ic,
lying
insec s
-
hey
a e
e icien
a
sampling
Lepidop e a,
lying
Coleop e a
and
T ichop e a,
bu
ca ch
ew
Nema oce a
and
no
non- lying
a h o-
pods.
Suc ion
ap
A
suc ion
ap
(Johnson,
1950;
Johnson
and
Taylo ,
1955)
consis s
o
a
an
which
d aws
ai
h ough
a
mesh
cone,
a
he
base
o
which
is
a
collec ing
cylinde .
Insec s
a e
d awn
in
wi h
he
ai
and
apped
in
he
cylinde ,
whe e
hey
a e
killed
by
a
py e h oid- ype
insec icide.
The
ca ch
can
be
sepa a ed
by
discs
which
a e
d opped
a
in e als
in o
he
cylinde .
Suc ion
aps
wo k
well
o
small,
ai bo ne
insec s
such
as
midges,
bu
a e
less
e icien
o
la ge
o
s ong- lying
ones
such
as
bee les
and
la ge
mo hs.
Malaise
ap
This
consis s
o
an
open- on ed
en
made
o
da k-colou ed
ne ing
(Sou hwood,
1966)
(Figu e
3.1).
The
oo
slopes
upwa ds
o
he
inne mos
co ne s,
whe e
a
small
hole
leads
o
a
collec ing
ube.
The
ap
is
usually
suspended
be ween
ees,
and
insec s
lying
in o
he
54
Fo aging
Beha iou
E ec s
o
ansmi e s
on
ba s
The
beha iou al
e ec s
due
o
dis u bance
caused
by
ca ching
ba s
and
a aching
he
ans-
mi e s
appea
o
be
minimal
-
En wis le
(1994)
compa ed
beha iou
o
P.
au i us
on
he
i s
nigh
a e
a achmen
and
on
subsequen
nigh s
and
ound
no
di e ence.
No
agged
ba s
abandoned
hei
oos s
in
he
s udy.
The
main
e ec
mus
be
ha
o
lying
wi h
a
hea y
weigh
a ached
and
an
ae ial
ailing
behind.
A
gene ally
assumed
ule
o
adio
ack-
ing
ba s
is
ha
hey
should
no
ca y
mo e
han
5%
o
hei
own
bodyweigh
(Ald idge
and
B igham,
1988).
En wis le
e
al.
(1996)
calcula ed
ha
ansmi e
mass
on
hei
P.
au i us
anged
om
6.4
o
9.5%
o
body
mass,
and
hose
o
Fuh mann
and
Sei z
(1992)
app oached
10%
in
mos
cases.
Adam
e
al.
(1994)
epo ed
ha
ansmi e s
we e
7.5%
o
body
weigh
o
male
C.
.
i ginianus
and
8.5%
o
emales
(since
males
we e
sligh ly
hea -
ie
han
he
pos -lac a ing
emales
in
hei
s udy).
They
obse ed
no
e ec
o
ca ying
ans-
mi e s
on adul s,
bu
ju eniles
appea ed
s essed
by
hem
and
agged
ones
did
no
lea e
he
oos
o
o age.
Hughes
and
Rayne
(1991)
conduc ed
a
s udy
on
cap i e
P.
au i us
in
a
ligh
oom
o
in es iga e
he
e ec s
o
ca ying
a i icial
loads.
They
poin ed
ou
ha
wing
loading
on
ba s
a ies
g ea ly
unde
na u al
condi ions
and
ha
ac o s
such
as
p e-hibe -
na ion
weigh
gain,
p egnancy,
weigh
o
mamma y
issue
and
he
load
o
a
ull
s omach
o
insec s
all
ha e
o
be
ca ied
in
ligh .
Also,
since
P.
au i us
has
a
low
wing
loading
and
b oad,
li -p oducing
wings,
i
should
be
capable
o
ca ying
hea ie
weigh s
han
o he
ba s
o
simila
size.
Hughes
and
Rayne
added
ishing
weigh s
be ween
he
shoulde
blades
o
hei
long-ea ed
ba s
and
obse ed
he
e ec s
in
ligh .
As
loading
inc eased,
he
ba s
lew
mo e
slowly
and
inc eased
hei
wingbea
equency.
Since
ae odynamic
heo y
p edic ed
ha
inc eased
loading
should
ha e
caused
he
ba s
o
ly
as e ,
his
was
unexpec ed.
They
concluded
ha
ca ying
ansmi e s
mus
ha e
an
e ec
on
ba s
as
small
as
P.
au i us.,
bu
ha u he
esea ch
was
needed.
En wis le
e
al.
(1996)
could
ind
no
ela ionship
be ween
ela i e
ansmi e
loading
( ansmi e
mass/body
mass)
and
ei he
he
u hes
dis ance
a elled
o
he ime
spen
close
o
he
oos
and
concluded
ha
he
ansmi e s
caused
no
impedimen
o
ligh .
Long-ea ed
ba s
a e
hus
ce ainly
capable
o
o aging
while
ca ying
ansmi e s,
and
any
ma ginal
e ec s
ha e
ye
o
be
shown.
ACTIVITY
PATTERNS
Eme gence
P.
au i us
eme ge
om
day
oos s
la e
in
he
e ening
ela i e
o
many
o he
espe ilionid
species.
En wis le
e
al.
(1996)
ound
ha
he
median
ime
o
eme gence
in
no h-eas
Sco land
was
55
min
a e
sunse
and
ha
eme gence
was
s ongly
co ela ed
wi h
bo h
ime
o
sunse
and
ime
o
ci il
wiligh
(de ined
as
he
pe iod
when
he
cen e
o
he
sun
is
be ween
0.8°
and
6.0°
below
he
ho izon).
In
B i ain,
pipis elles
eme ge
35
min
a e
sun-
se
(Swi ,
1980)
and
la ge,
as - lying
species
such
as
se o ines
(Ep esicus
se o inus;
Ca o
e
al.,
1995)
and
noc ules
(Nyc alus
noc ula;
Racey,
1991)
e en
ea lie
(see
also
Rydell
e
al.,
1996).
Jones
and
Rydell
(1994)
calcula ed,
om
all
a ailable
da a
om
Eu ope,
ha
he
a e age
median
ime
o
eme gence
was
44
min
a e
sunse
in
P.
au i us,
compa ed
wi h
33
min
in
P.
pipis ellus,
8
min
in
N.
noc ula
and
11
min
in
E.
se o inus.
The
ime
o
eme gence
o
Co yno hinus
ownsendii
ingens,
a
close
Ame ican
ela i e
o
Pleco us
species,
was
ound
Ac i i y
Pa e ns
55
o
be
45.5
min
a e
sunse
(Cla k
e
al.,
1993).
Ac i i y
pa e ns
in
insec i o ous
ba s
a e
gene ally
conside ed
o
be
con olled
by
an
endogenous
ci cadian
hy hm
synch onized
o
he
daily
ligh
cycle
by
ligh -sampling
beha iou
(de
Cou sey
and
de
Cou sey,
1964),
and
indi iduals
eme ge
wi hin
e y
na ow
bands
o ligh
in ensi y
(E ke
and
K ach ,
1978).
The
high
deg ee
o
co ela ion
be ween
ime
o
eme gence
and
he
imes
o
sunse
and
ci il
wiligh
indica e
ha
P.
au i us
is
no
excep ion.
The
leng h
o
ci il
wiligh
inc eases
wi h
la i ude
in
summe
(Howa d,
1995),
and
hus
he
ime
o
eme gence
may
be
expec ed
o
be
la e
ela i e
o
sunse
u he
no h
in
he
ba s'
ange.
This
does
appea
o
be
he
case
(Table
4.1);
al hough
he
ou
s udies
lis ed
in
he
able
used
sligh ly
di e en
me hods o
es ima e
when
ba s
eme ged,
he e
is
a
s ongly
linea
ela ionship
be ween
ime
o
eme -
gence
and
la i ude
(
=
0.96;/><0.01),
indica ing
ha
long-ea ed
ba s
eme ge
ela i ely
la e
u he
no h,
whe e
wiligh
las s
longe .
Howa d
(1995)
calcula ed
ha
ci il
wiligh
las s
a ound
75
minu es
a
58°N
and
a ound
45
minu es
a
50°N
in
midsumme .
P.
au i us
hus
eme ged
10-19
min
be o e
he
end
o
ci il
wiligh
(Table
4.1),
a
which
ime
ligh
in en-
si y
is
below
abou
3
lux
(Swi ,
1997).
Mean
ligh
in ensi y
a
57°N
was
measu ed a
0.7
lux
1
h
a e
sunse
(Rydell
e
al.,
1996).
The
la e
eme gence
o
b own
long-ea ed
ba s
is
almos
ce ainly
connec ed
wi h
hei
habi
o
gleaning.
Time
o
eme gence
in
any
insec i o ous
ba
is
likely
o
be
a
comp omise
be ween
he
need
o
go
ou
and
eed
and
he
isk
o
p eda ion
a
high
ligh
le els
(Speakman,
1991a; Jones
and
Rydell,
1994).
P eda ion
by
bi ds,
mainly
kes els
and
awny,
ba n
and
long-ea ed
owls,
has
been
es ima ed
o
accoun
o
abou
11%
o
he
annual
mo -
ali y
o
B i ish
ba s
(Speakman,
1991a;
see
also
Chap e
7).
Speakman
(1991a)
concluded
ha ,
al hough
ba s
comp ise
only
a
small
p opo ion
o
he
p ey
aken
by
hei
p eda o s,
he
e ec
o
he
p essu e
his
p eda ion
has
on
ba
beha iou
canno
be
igno ed.
Species
such
as
pipis elles,
which
a e
ae ial
hun e s
and
ely
o
mos
o
hei
ood
on
he
dusk
and
dawn
peaks
o
lying
insec s
(Swi ,
1980),
ha e
o
eme ge
a
ela i ely
high
ligh
le els
and
isk
being
caugh
by
p eda o s
bu ,
because
P.
au i us
gleans
much
o
i s
p ey,
i
elies
a
TABLE
4.1
Times
o
eme gence
o
P.
au i us
a
di e en
la i udes
in
Eu ope.
S udy
La i ude
A e age
in e al
be ween
sunse
and
eme gence
(min)
No e
En wis le
e
al.
(1996)
57°N
55
Median
eme gence;
adio- acked
ba s
Howa d
(1995)
51°23'N
40
Calcula ed
a e age
ime
o
eme gence
om
oos
coun s
Fuh mann
and
Sei z
(1992)
50°N
26
A e age
ime
o
eme -
gence
o
indi idual
adio- acked
ba s;
ange
=
7—43
min
Ba a aud
(1990)
46°N
15
Time
o
a i al
o
ma ked
ba s
a
eeding
si es

56
Fo aging
Beha iou
less
on
his
dusk
peak.
Non- lying
p ey
a e
a ailable
all
nigh ,
and
a
u he
educ ion
in
he
p essu e
o
ea ly
eme gence
occu s
because
he
peak
ac i i y
o
mo hs
occu s
a ound
mid-
nigh
(Rydell
e al.,
1996).
Ea ly
eme gence
would
hus
ca y
no
bene i s
o
long-ea ed
ba s
and
would
inc ease
hei
isk
o
p eda ion,
which
in
any
case
is
high
because
hey
ly
slowly.
La e
eme gence
is
co ela ed
wi h
gleaning
in
many
ba
species
in
empe a e
egions
(Jones
and
Rydell,
1994)
and
his
may
no
only
be
because
o
ood
a ailabili y.
I
appea s
ha
gleaning
i sel
is
a
dange ous
occupa ion
unless
i
is
done
in
he
da k,
since
gleane s
ha e
o
ly
slowly.
Taake
(1985)
ound
ha ,
o
a
sample
o
Eu opean
ba
species,
hose
wi h
pale
colou ed
en al
u
eme ged
la e
in
he
e ening
and
ended
o
be
gleane s.
O he
Eu opean
species
which
glean
a
leas
pa
o
hei
die
also
eme ge
la e
in
he
e ening
(e.g.
Myo is
na e e i;
Swi ,
1997),
as
do
closely
ela ed
Ame ican
species
such
as
Co yno hinus
ownsendii
(Cla k
e
al.,
1993).
Eme gence
Beha iou
In
some
espe ilionid
species
(e.g.
P.
pipis ellus-,
Swi ,
1980;
Bullock
e
al.,
1987),
ou -
bu s ,
o
clus e ing,
beha iou
has
been
desc ibed,
in
which
pe iods
when
la ge
numbe s
o
ba s
eme ge
a e
in e spe sed
wi h
pe iods
when
ew,
i
any,
lea e
he
oos .
No
de ailed
s udies
on
he
eme gence
o
long-ea ed
ba s
ha e
been
conduc ed,
bu
anecdo al
obse a-
ions
indica e
ha
hey
do
some imes
eme ge
in
g oups
o
wo
o
h ee
and
ha
he
eme -
gence
o
one
equen ly
seems
o
lead
o
he
eme gence
o
o he s.
Howe e ,
no
de ini e
clumping,
o
ou bu s
beha iou ,
such
as
is
ob ious
in
pipis elles,
can
be
obse ed.
Speakman
e
al.
(1992)
ound
ha
signi ican
clus e ing
was
a
ea u e
only
o
la ge
(>
150)
colonies
o pipis elles
and
ha
a
colony
size
o
less
han
100,
eme gence
was
andom.
They
sugges ed
ha
clus e ing
may
be
due
ei he
o
p eda o
a oiding
beha iou
o
o
he
e ec
o
p essu e
on
exi
holes,
and
poin ed
ou
ha ,
while
p eda o
a oidance
may
be
an
e ec
o
clumped
eme gence
in
la ge
colonies,
i
may
no
be
he
main
eason
o
he
beha -
iou .
P.
au i us
o m
small
colonies
(Chap e
6),
mos
o
which
numbe
ewe
han
80
indi-
iduals,
and
he e o e
hey
a e
p obably
ne e la ge
enough
o
p essu e
on
exi
holes
o
become
an
issue.
Thus,
al hough
he
obse ed
depa u e
om
he
oos
o
small
g oups
o
ba s
may
be
a
mild
o m
o
an i-p eda o ,
o
'sa e y
in
numbe s'
beha iou ,
p ope
clumped,
o
ou bu s ,
eme gence
does
no
occu .
The
ime
aken
o
a
colony
o
eme ge
has
been
epo ed
o
be
independen
o
colony
size
(Howa d,
1995),
again
p obably
due
o
he
o ma ion
o
small
colonies
by
his
species
and
consequen
lack
o
p essu e
on
exi
holes.
Ligh -sampling
beha iou , whe eby
ba s
check
he
le el
o
ligh
in ensi y
as
a
means
o
synch onizing
hei
ac i i y
hy hm
o
he
daily
ligh
cycle
(de
Cou sey
and
de
Cou sey,
1964;
Vou e
e
al.,
1974),
has
been
obse ed
inside
P.
au i us
oos s
(Swi ,
1981).
In
he
10-15
min
be o e
eme gence,
ba s
lew
inside he
oos ,
and
du ing
ligh s
hey
we e
seen
o
ly
low
o e
he
exi
hole,
be o e
aligh ing
b ie ly
on
oo
beams
and hen
epea ing
he
beha iou .
Finally,
hey
aligh ed
beside
he
exi
hole
o
a
ew
seconds
and hen
d opped
h ough
i .
E ec s
o
En i onmen al
Fac o s
on
Eme gence
Rain
du ing
he
expec ed
pe iod
o
eme gence
was
ound
signi ican ly
o
delay
he
ime
o
eme gence,
bu
low
ai
empe a u es
(<7°C)
on
d y
nigh s
did
no
p e en
ba s
om
Ac i i y
Pa e ns
57
eme ging
(En wis le
e al.,
1996).
Hea y
cloud
co e
esul ed
in
ea lie
han
no mal
eme -
gence
(Howa d,
1995),
bu
ligh
cloud
had
li le
e ec .
The
a e age
ime
o
eme gence
was
ound
o
a y
signi ican ly
be ween
oos s
wi hin
an
a ea
and
o
be
s ongly
co ela ed
wi h
he
dis ance
om
he
oos
o
he
closes
woodland
(En wis le
e
al.,
1996).
Ba s
eme ged
ea lie
a
oos s
close
o
woodland
(Figu e
4.2),
a
inding
simila
o
ha
o
Jones
e
al.
(1995)
o
Rhinolophus
e umequinum-,
hese
au ho s
sugges ed
ha
woodland
loca ed
close
o
oos s
may
p o ide
co e
om
ae ial
p eda o s
and
so
enable
ba s
o
eme ge
a
el-
a i ely
high
ligh
le els.
The
use
o
oos s
close
o
woodland hus
allows
long-ea ed
ba s
o
ex end
hei
o aging
pe iod
and
is
an
impo an
conside a ion
o
hei
conse a ion.
I
has
been
sugges ed
as
one
o
he
easons
why
P.
au i us
selec
oos s
close
o
woodland
(En wis le
e al.,
1997).
Eme gence
in
P.
aus iacus
Da a
a e
spa se
on
imes
o
eme gence
in
his
species.
Baue o a
(1982)
epo ed
i
eme ged
a ound
dusk
(possibly
a ound
he
end
o
ci il
wiligh ).
Ba a aud
(1990)
obse ed
ha
ligh - agged
specimens
o
bo h
P.
au i us
and
P.
aus iacus
a i ed
a
o aging
a eas
app oxi-
ma ely
15
min
a e
sunse .
A
single
indi idual
adio- acked
by
Fluckige
and
Beck
(1995)
le :
he
oos
abou
hal
an
hou
a e
sunse .
The
ime
o
eme gence
hus
appea s
o
be
80
Dis ance
o
woodland
(km)
FIG
4.2
A e age
ime
o
eme gence
o
adio- acked
P.
au i us
om
six
oos s,
plo -
ed
agains
he
dis ance
om
each
oos
o
he
nea es
woodland
( ep oduced
wi h
pe -
mission
om
En wis le
e
al.,
1996).
2
=
0.79;
p
<
0.02.
58
Fo aging
Beha iou
simila
o
ha
o
P.
au i us
and
in
iew
o
hei
b oadly
simila
die s
and
o aging
habi s,
his
seems
o
be
easonable.
The
e ec s
o
en i onmen al
ac o s
on
eme gence
ha e
no
ye
been
documen ed.
Ac i i y
In
bo h
P.
au i us
and
P.
aus iacus
(on
which,
again,
limi ed
da a
a e
a ailable),
he
nigh ly
ac i i y
pa e n
has
been
shown
o
be
unimodal
(Baue o a,
1982;
Fuh mann
and
Sei z,
1992;
Fluckige
and
Beck,
1995;
En wis le
e
al.,
1996).
A e
eme gence,
ba s
emain
ac i e
ou side
he
oos
all
nigh .
En wis le
e
al.
(1996)
ound
ha
P.
au i us
in
Sco land
e u ned
o
he
oos
a
median
o
57
min
be o e
sun ise.
Like
eme gence,
he ime
o
e u n
was
signi ican ly
ela ed
o
bo h
ime
o
sun ise
and
ime
o
ci il
wiligh .
In
Ge many,
whe e
he
nigh s
a e
longe
in
summe ,
he
ime
o
e u n
o
he
oos
o
his
species
was
ea lie ,
a
an
a e age
o
190
min
be o e
sun ise,
and
ba s
did
no
lea e
again
be o e
dawn
(Fuh mann
and
Sei z,
1992).
In
bo h
s udies,
ac i i y
was
cha ac e ized
by
pe iods
o
ligh
in e spe sed
wi h
sho
pe iods
o
inac i i y
which
anged
om
1
o
65
min
in
du a ion.
P.
aus iacus
showed
simila
beha iou
(Fluckige
and
Beck,
1995);
an
indi idual
was
ac i e
o
mos
o
he
nigh
and
emained
s a iona y
only
occasionally.
The
unimodal
ac i i y
pa e n
o
long-ea ed
ba s
con as s
wi h
hose
o
many
o he
em-
pe a e
zone
ba
species,
which
a e
bimodal
wi h
peaks
a ound
dusk
and
dawn
(E ke ,
1982).
Such
bimodal
pa e ns
a e
he
commones
ype
o daily
hy hms
among
animals
(Ascho ,
1966),
wi h
he
ough
o
ac i i y
occu ing
a ound
midday
o
midnigh ,
and
wi h
he
second
peak
being
lowe
and
equen ly
educed
i
en i onmen al
s imuli
a e
emo ed.
In
some
ba
species
(e.g.
P.
pipis ellus-,
Swi ,
1980;
Maie ,
1992),
he
dawn
peak
may
be
supp essed
a
ce ain
imes
o
yea
and
may
only
be
ob ious
du ing
lac a ion,
when
he
high
ene gy
demands
on
emales
cause
hem
o
o age
wi h
maximum
e iciency.
A
hese
imes,
he
dusk
and
dawn
peaks,
which
coincide
wi h
he
peaks
in
ac i i y
shown
by
hei
lying
insec
p ey,
a e
sepa a ed
by
a
pe iod
o
nigh
oos ing,
a
which
ime
he
young
a e
suckled.
The
eason
o
he
unusual
unimodal
pa e n
in
long-ea ed
ba s
is
p obably
p ey
a ail-
abili y.
Mo hs
a e
unlike
mos
lying
insec s
in
ha
hey
a e
ac i e
all
nigh ,
wi h
hei
ac i -
i y
peak
a ound
midnigh
(Rydell
e
al.,
1996);
Pleco us
a e
he e o e
able
o
keep
o aging
p o i ably
a
a
ime
when
non-mo h
eede s
canno
do
so.
Non- lying
a h opods,
es ing
mo hs
and
diu nal
insec s
a e
also
equally
a ailable
all
nigh
and
hus
gleaning,
oo,
is
p o i able
in
he
middle
o
he
nigh .
Long-ea ed
ba s
eme ge
la e
and
keep
o aging
un il
hey
ha e
sa is ied
hei
ene gy
equi emen s,
a e
which
hey
e u n
o
he
oos
and
s ay
he e.
A
high
la i ude,
indi iduals
may
o age
almos
con inuously
du ing
he
hou s o
da kness
(En wis le
e
al.,
1996),
while
u he
sou h
hey
ha e
been
ound
o
e u n
o
hei
day
oos s
up
o
573
min
be o e
sun ise
and
no
o
lea e
again.
En wis le
e
al.
(1996)
ound
ha
he
only
ba s
which
e u ned
o
he
oos
du ing
he
nigh
we e
lac a ing
emales.
When
he
young
we e
newly
bo n,
mo he s
made
se e al
is-
i s
o
sho
du a ion,
while
la e
in
he
summe
hey
en e ed
he
oos
less
o en
bu
s ayed
longe .
The
eason
o
hese
e u ns
was
almos
ce ainly
o
suckle
in an s.
Unlike
many
oden s,
which
suckle
hei
young
in equen ly,
i
appea s
ha
equen
suckling
is
he
no m
in
insec i o ous
ba s.
Wilde
e
al.
(1995)
sugges ed
ha ,
because
a
mo he 's
me a-
bolic
a e
du ing
ligh
is
maximal,
milk
p oduc ion
is
also
apid
du ing
o aging
and
Fo aging
59
accumula ion
o
milk
in
he
mamma y
glands
means
emales
ha e
o
eed
hei
young
se e al
imes
pe
nigh .
F equen
suckling
also
elie es
au oc ine
eedback
and
allows
maximal
milk
p oduc ion
du ing
subsequen
bou s
o
o aging.
Two
sub-species
o
Co yno hinus
ownsendii,
C.
.
i ginianus
and
C.
.
ingens,
we e
bo h
ound
o
ha e
ac i i y
pa e ns
e y
simila
o
P.
au i us
(Bagley
and
Jacobs,
1985;
Cla k
e
al.,
1993).
Bo h
ha
unimodal
pa e ns,
wi h
lac a ing
emales
making
up
o
h ee
e u ns
each
nigh
o
he
ma e ni y
oos .
I
seems
ha
emales
o
ba
species
wi h
bimodal
ac i i y
pa e ns
combine
a
e u n
o
he
oos
o
suckle
in an s
wi h
hei
ac i i y
ough
in
he
middle
o
he
nigh ,
bu
long-ea ed
ba s
and
hei
ela i es
ha e
o
inco po a e
isi s
o
hei
in an s
in o
hei
main
pe iod
o
ac i i y.
Howa d
(1995)
se
up
an
au oma ic
eco ding
sys em
based
on
an
in a ed
beam
o
moni o
ac i i y
inside
a
P.
au i us
oos .
His
esul s
show
ha ,
while
ac i i y
a
nigh
was
highes
du ing
lac a ion,
he e
was
also
some
ac i i y
inside
he
oos
a
o he
imes
o
yea ,
indica ing
ha
some
non-lac a ing
ba s
did
e u n
du ing
he
nigh .
He
sugges ed
ha
ba s
which
o aged
close
o
he
oos
may
ha e
e u ned
o
sho
es ing
pe iods.
An
al e na i e
explana ion
is
ha
he
eco ded
ac i i y
was
due
o
some
ba s
cu -
ailing
hei
o aging
ea ly,
as
was
es ablished
in
Fuh mann
and
Sei z
s
(1992)
s udy.
E ec s
o
En i onmen al
Fac o s
on
Ac i i y
Rain
inhibi ed
ac i i y
away
om
he
oos
as
well
as
p e en ing
eme gence
(En wis le
e
al.,
1996);
ba s
hung
up
and
became
inac i e
immedia ely
ain
s a ed
and
he e
was
a
sig-
ni ican
inc ease
in
hanging
ime
du ing
we
nigh s.
These
au ho s
sugges ed
ha
he
appa en
eluc ance
o
ly
in
ain
(sha ed
by
mos
ba
species;
E ke ,
1982)
is
mos
p ob-
ably
due
o
p oblems
linked
wi h
he mo egula ion
-
we
ba s
lose
hea
apidly.
Al e na i ely,
hey
sugges ed,
he
sound
o
alling
ain
may
in e e e
wi h
he
ba s
sys em
o
passi e
lis ening
o
p ey.
Ai
empe a u es
as
low
as
3.5°C
did
no
inhibi
o aging
by
P.
au i us
(En wis le
e
al.,
1996),
unlike
he
si ua ion
in
species
which
eed
on
lying
insec s
-
al hough
low
ai
empe a u es
may
ha e
inhibi ed
insec
ligh ,
es ing
insec s
we e
s ill
a ailable
o
gleaning.
Gleaning
may
also
make
mis
and
og
less
o
a
p oblem
o
long-
ea ed
ba s
han
o
exclusi ely
ae ial
o aging
species.
Mis d ople s
in
he
a mosphe e
abso b
ul asound
(Sales
and
Pye,
1974)
and
so
cause
a
educ ion
in
he
hun ing
e iciency
o
ba s
such
as
pipis elles
(Swi ,
1980),
bu
may
no
hampe
passi e
lis ening
o
he
same
ex en .
Howe e ,
adio- agged
P.
au i us
which
eme ged
on
oggy
nigh s
we e
ound
o
show
abno mal
ligh
pa e ns
and
e en
some imes
o
cu ail
o aging
(A.C. En wis le,
pe s.
comm.),
indica ing
ha
mis
and
og
do
ad e sely
a ec
hei
o aging
o
some
deg ee.
The
e ec
o
wind
is
likely
o
be
simila
o
ha
on
mos
ba
species
-
on
windy
nigh s
indi id-
uals
o age
mo e
in
shel e ed
a eas
such
as
woodland
and
behind
hedges.
Insec s
accumu-
la e
in
such
shel e ed
habi a s
unde
hese
condi ions.
FORAGING
Flyways
Bo h
P.
au i us
and
P.
aus iacus
consis en ly
use
landscape
ea u es
such
as
hedges
o
ee-
lines
along
which
o
ly
be ween
oos s
and
o aging
a eas
and
be ween
eeding
si es.
60
Fo aging
Beha iou
Fea u es
epo ed
o
be
used
a e
usually
linea
and,
besides
hedges
and
eelines,
include
o e g own
banks,
ences,
o es
ides,
ailway
lines
and
s eams
wi h
ege a ion
on
he
banks.
Indi idual
ba s
use
he
same
lyways
h oughou
he
summe
(Ba a aud,
1990),
and
H.gh
along
hem
is
epo ed
o
be
close
o
he
g ound
and
as e
han
no mal
o aging
ligh
(Ba a aud,
1990;
En wisde
e al.,
1996).
Howa d
(1995)
measu ed
he
speed
a
which
P.
au i us
lew
du ing
commu ing
ligh s
along
a
lyway
and
es ima ed
hei
speed
o
be
almos
mi!i
'
Is
L
COn Slde abIy
as e
han
hei
ligh
du ing
o aging
(Chap e
2)
and
com-
pa able
wi h
ha
o
o he
espe ihonid
species
o
simila
size.
Howa d's
obse a ions
o
P.
au i us
along
lyways
showed
ha
hey
ne e
c ossed
open
spaces,
e en
i
using
he
lyways
in ol ed
co e ing conside able
ex a
dis ances.
When
commu ing
o
a
o aging
a ea
in
a
plan a ion
200
m
om
he
oos ac oss
an
open
ield,
ba s
ne e
lew
o e
he
ield,
bu
a -
elled
abou
300
m
o
he
plan a ion
by
way
o
he
hedge
a ound
he
edge.
En wis le
(1994)
simila ly
epo ed
ha
P.
au i us
co e ed
ex a
dis ances
because
o
hei
use
o
lyways.
Two
possible
easons
o
he
use
o
lyways
a e
o
educe
he
isk
o
ae ial
p eda ion
o
as
na iga ional
aids.
Al hough
bo h
long-ea ed
species
do
occasionally
ca ch
p ey
while
lying
a
ong
hedges
(Ba a aud,
1990),
he
ba s'
as ,
s aigh
ligh
sugges s
his
is
no
hei
main
pu pose.
The
possibili y
o
p eda ion
by
bi ds
is
p obably
a
se ious
enough h ea
o
p e-
en
ela i ely
slow- lying
ba s
(e en
i
hey
a e
lying
a
wha
is,
o
hem,
a
as
speed)
om
yin|un
L
°pen"
AJ e na l ely>
11
is
possible
ha ,
because
he
echoloca ion
calls
o
long-
ea ed
ba s
ha e
eiy
limi ed
ange,
hey
ha e
o
ely
on
landscape
ea u es
as
na iga ional
aids,
as
sugges ed
by
Howa d
(1995).
Flyways
a e
egula ly
used
by
o he
ba s
wi h
low-
in ensi y
calls,
such
asMyo is
na e e i
(Swi ,
1997),
in e ing
ha
he
na iga ion
explana-
ion
is
mo e
likely.
On
he
o he
hand,
M.
na e e i
also
lies
slowly
and
mus
be
as
ulne able
as
Pleco us
o
p eda ion.
Myo is
dauben onii
use
lyways
and
ne e
c oss
open
g ound, and
hei
echoloca ion
calls
a e
much mo e
in ense.
In
any
case,
he
wo
explana-
ions
a e
no
mu ually
exclusi e
and
i
is
qui e
p obable
ha
long-ea ed
ba s
use
lyways
o
bo h
he
abo e
easons.
Eckman
and
de
Jong
(1996)
ound
ha
P.
au i us
in
Sweden
a oided
o aging
in
isola ed
pa ches
o
o es
in
an
ag icul u al
a ea
o
o e
o es ed
islands
in
a
shallow
lake.
They
a ibu ed
his
a oidance
o
he
isola ion
a he
han
o
insec
abun-
ance,
which
was
ac ually
highe
in
he
open
a eas,
and
sugges ed
ha
p eda ion
isk
p e-
en ed
ba s
om
c ossing
open
ields
o
wa e
o
each
p oduc i e
o aging
a eas.
This
s udy
emphasized
he
impo ance
o
lyways
o
P.
au i us
and
he
nega i e
impac
o
habi-
a
pa chiness
o
his
species.
Fo aging
Habi a
P.
au i us
His o ically,
his
species
has
always
been
desc ibed
as
a
woodland
o age
and
has
been
con-
side ed
as
a
ypical
oliage
gleane
in
deciduous
o es s.
Howe e ,
jus
as
i
has
been
shown
(Chap e
3)
o
be
a
mo e
adap able
han
his
in
i s
die ,
so
mo e
ecen
s udies
ha e
indi-
ca ed
ha
a
a ie y
o
al e na i e
habi a s
may
also
be
used.
Swi
and
Racey
(1983)
obse ed
P.
au i us
ma ked
wi h
e lec i e
ings
o aging
in
deciduous
woodland,
bi ch
sc ub
and
a
ga den
con aining
many
ma u e
coni e ous
and
deciduous
ees,
and
Ba a aud
(1990)
mis -ne ed
indi iduals
in
illage
ga dens,
sca e ed
woodlands,
o cha ds
and
pa k-
land
among
meadows.
He
also
emphasized
he
impo ance
o
ag icul u al
buildings
as
o -
aging
si es
-
ba s
pu sued
insec s
in o
s ables
and
sheep olds
h ough
open
doo s
and
also
Fo aging
61
gleaned
mo hs
a
es
inside.
They
hun ed
ex ensi ely
inside
he
buildings
du ing
we
wea he ,
an
oppo unis ic
mo e
which
enabled
hem
o
keep
o aging
when
condi ions
ou -
side
we e
unsui able.
Heise
and
Schmid
(1988)
obse ed
b own
long-ea ed
ba s
o aging
in
a eas
con aining
low
bushes,
as
well
as
in
o es s.
Two
we e
caugh
in
mis -ne s
only
50
cm
abo e
he
g ound,
indica ing
ha
hey
we e
p obably
gleaning
om
hese
bushes.
The e
is
also
e idence
om
insec s
consumed
(Robinson,
1990;
Shiel
e
al.,
1991)
ha
he
species
may
glean
om
g assland
and
pas u e
a
imes.
Quan i a i e
da a
on
he
ela i e
impo ance
o
a ious
o aging
habi a s
ha e
been
p o-
ided
by
wo
ecen
s udies
using
adio
acking.
The
i s
o
hese
was
by
Fuh mann
and
Sei z
(1992)
in
Ge many.
Six
emales
(all
non- ep oduc i e
o
lac a ing)
we e
acked
and
hey
spen
mos
o
hei
ime
in
woodland.
O
he
o al
o aging
ime
eco ded,
55-0%
was
spen
in woodland
wi h
widely
spaced
ees
(mo e
han
2
m
be ween
unks)
and
29.3%
in
woodland wi h
ees
plan ed
close
oge he
(less
han
2
m
be ween
unks).
The
emaining
ime
(15.7%)
was
spen
in
ga dens,
pa kland,
o cha ds
o
along
ailway
lines
o
mo o way
embankmen s.
En wis le
e
al.
(1996)
acked
16
ba s
o e
a
o al
o
65
nigh s
in
no h-eas
Sco land.
All
o
he
75
eeding
si es
loca ed
we e
associa ed
ei he
wi h woodland
o
wi h
indi idual
ees;
17%
we e
in
ees
adjoining
pas u e
and
only
9%
ound
ees
close
o
wa e .
Ba s
spen
42%
o
he
o al
o aging
ime
in
ma u e
deciduous
woodland
(excluding
bi ch
woodland),
and
when
using
non-na i e
coni e ous
plan a ions
(mainly
sp uce)
hey
emained
on
he
edge
and
did
no
use
he
inne
woodland.
The
ela i e
ime
spen
in
di -
e en
ypes
o
woodland
is
shown
in
Figu e
4.3.
To
in es iga e
whe he
ba s
used
deciduous
woodland
p e e en ially,
En wis le
e
al.
(1996)
pe o med
a
u iliza ion—a ailabili y
es
based
on
ime
budge s.
They
es ima ed
he
p opo ion
o
a ious
ypes
o
woodland
o
g oups
o
ees
wi hin
o aging
ange
o
he
oos
om
la ge-scale
maps
and
hen
compa ed
his
wi h
he ime
ba s
ac ually
spen
in
each
ype
by
a
Wilcoxon
pai ed
es .
As
shown
by
Figu e
4.4,
he
ba s
spen
signi ican ly
mo e
ime
in
ma u e
deciduous
woodland
han
would
be
expec ed
i
hei
use
o
all
ees
was
andom,
i.e.
hey
we e
selec ing
his
ype
o
woodland.
Sweep
ne
samples
in
o aging
a eas
showed
ha ,
while
he e
we e
no
di e ences
in
he
o al
numbe s
o
insec s
a ailable
be ween
woodland
ypes,
ma u e
deciduous
woodland
did
p oduce
signi ican ly
highe
numbe s
o
mo hs.
The
au ho s
sugges ed
ha
his
was
he
eason
o
he
ba s
o aging
p e e en ially
he e,
since
o he
ac o s
impo an
o
he
ba s,
he
p o ision
o
su aces
om
which
insec s
could
be
gleaned
and
co e
om
ae ial
p eda ion,
we e
p o ided
equally
by
all
ypes
o
woodland.
Mo hs
a e
he
p e e ed
ood
o
P.
au i us,
so
hey
o age
p e e en-
ially
in
he
habi a s
in
which
mos
mo hs
a e
o
be
ound.
Jus
as
hei
die
is
lexible,
how-
e e ,
so
ba s
o
his
species
a e
also
able
o
adap
hei
o aging
beha iou
o
include
o he
habi a s
con aining
ees
and
bushes.
P.
aus iacus
Fo aging
habi a s
o
his
species
a e
epo ed
o
include
woodland
(Ba a aud,
1990),
as
well
as
open
meadows
(Baue o a,
1982)
and
pas u e
con aining
ees
(D.
La oley,pe s.
comm.).
A
single
non-lac a ing
emale
was
adio
acked
o e
ou
nigh s
by
Fluckige
and
Beck
(1995).
This
ba
o aged
in
six
dis inc
a eas,
including
ound
a
ches nu
ee
in
a
ga den
close
o
he
oos ,
open
meadows,
a
che y
o cha d,
o es
edge
and
mixed
woodland.
I
hus
di e s
om
P.
au i us
in
hun ing
in
open
habi a s
such
as
meadows,
and
Fluckige
and
Beck
(1995)
epo ed
ha
i
seemed
o
ha e
wo
sepa a e
o aging
s a egies,
one
o
62
Fo aging
Beha iou
Woodland
ype
FIG
4.3
The
pe cen age
o
ligh
ime
o
16
adio- acked
P.
au i us
spen
wi hin
di e en
ypes
o
woodland
in
no h-eas
Sco land,
n
=
65
ba
nigh s
plus
7
hal
nigh s
( ep oduced
wi h
pe mission
om
En wis le
e
al.,
1996).
clu e ed
en i onmen s
which
included
gleaning
and
he
o he ,
in ol ing
as e
ligh ,
o
open
si ua ions.
This
explains
why
he e
a e
ewe
gleaned p ey
i ems
in
he
die
o
P.
aus-
iacus
han
in
ha
o
P.
au i us.
P.
aus iacus
also
di e s
om
P.
au i us
in
ha
i
is
epo ed
o
ca ch
mo hs
in
ligh
ound
s ee
lamps
(see
Chap e
9).
Use
o
Feeding
Si es
Bo h
b own
and
g ey
long-ea ed
ba s
ha e
been
epo ed
o
use
a
se ies
o
eeding
si es
du -
ing
a
nigh 's
o aging,
and
o
ly
be ween
si es
quickly
and
di ec ly,
using
lyways.
P.
au i us
Fligh
wi hin
si es
is
slow
and
dipping
(En wis le
e
al.,
1996;
Swi
and
Racey,
1983),
and
ba s
o en
ly
among
he
b anches
o
ees.
The
ime
spen
in
any
one
a ea
is
e y
a iable
Fo aging
63
1.0
0.8
0.6
0.4
0.2
_J
I I
'—
0.2
0.4
0.6
0.8
P opo ion
o
woodland
deciduous
1.0
FIG
4.4
U iliza ion-a ailabili y
es
o
compa e
he
ime
spen
by
indi idual
?W/ -
acked
P.
au i us
in
deciduous
woodland
wi h
he
p opo ion
o
a ailable
woodland
which
was
deciduous.
The
line
ep esen s
he
ime
each
ba
would
spend
in
deciduous
woodland
i
i s
use we e
andom
( ep oduced
wi h
pe mission
om
En wis le
e
al.,
1996).
(Fuh mann
and
Sei z,
1992),
bu
s ays
o
up
o
30
min
(Swi
and
Racey,
1983)
and
33
min
(Ba a aud,
1990)
ha e
been
eco ded.
The
a e age
size
o
ligh
a eas
is
also
a iable,
and
anged om
0.3
o
10.5
ha
in
Fuh mann
and
Sei z's
s udy.
En wis le
e
al.
(1996)
eco ded
ha
indi idual
ba s
used
be ween
1
and
9
(median
=
3)
eeding
si es
pe
nigh
and
he
num-
be
o
ligh s
be ween
si es
anged
om
1
o
16
(median
=
6).
They
also
ound
a
high
nigh -
o-nigh
p edic abili y
as
o
which
si es
a
pa icula
indi idual
would
use
-
each
used
a
1
i
mi ed
numbe
a
any
ime
o
yea
and
o
all
si es,
77%
we e
used
by
he
same
ba
on
mo e
han
one
nigh .
Females
we e
signi ican ly
mo e
likely
o
use
he
same
si e
om
nigh
o
nigh
han
we e
males,
which
hus
appea ed
o
be
mo e
oppo unis ic
in
hei
choice
o
si es.
Si es
o
which
emales
e u ned
we e
hose
which
had
been
used
o
longe
he
nigh
be o e,
indi-
ca ing
ha
ba s
emembe ed
which
si es
we e
mos
p oduc i e
and
e u ned
o
hem.
Si es
we e
equen ly
used
by
wo
o
mo e
ba s
a
he
same
ime
(an
obse a ion
also
made
by
Fuh mann
and
Sei z,
1992).
Up
o
i e
ba s
we e
eco ded
eeding
oge he ,
wi h
no
ob i-
ous
in e ac ions
be ween
hem.
Da a
showed
ha
a
ba
was
signi ican ly
mo e
likely
o
use
64
Fo aging
Beha iou
a
si e
i
ano he
indi idual
was
also
p esen .
In
some
species,
such
beha iou
is
hough
o
be
due
o
social
acili a ion
(Wilkinson,
1995),
whe eby
ba s a e
a ac ed
o
ich
ood
pa ches
by
he
echoloca ion
calls
o
conspeci ics.
In
his
way,
hey
bene i
om
he
o aging
success
o
o he
indi iduals.
Howe e ,
because
o
he
e y
sho
ange
o
Pleco us
calls,
En wis le
e
al.
(1996)
conside ed
i
unlikely
ha
his
was
he
case
in
hei
s udy,
and
i
was
mo e
p obable
ha
indi iduals
ound
p oduc i e
eeding
si es
independen ly.
They
ound
no
e idence
o
agg essi e
in e ac ions
be ween
P.
au i us
o aging
oge he
-
a
simila
si u-
a ion
o
ha
epo ed
by
Racey
and
Swi
(1985)
o
pipis elles
in
ich
ood
pa ches.
P.
aus iacus
Fluckige
and
Beck
(1995)
ound
ha
his
species
simila ly
used
si es
on
successi e
nigh s
and
also
e- isi ed
a
si e
on
he
same
nigh
on
occasions.
Cas o
e
al.
(1993)
epo ed
ha
g ey
long-ea ed
ba s
used
eeding
si es
oppo unis ically,
a ying
si es
acco ding
o
wea he
condi ions
and
ood
a ailabili y,
and
Baue o a
(1982)
also
ound
hem
o
be
oppo unis ic
in
hei
use
o
s ee
lamps.
Indi iduals
lew
as
and
s aigh
unde
a
line
o
s ee
lamps,
pu suing
la ge
mo hs
which
had
been
a ac ed
o
he
ligh .
Fas ,
s aigh
ligh
was
also
used
when
hun ing
abo e
meadows,
bu
in
ga dens
and
woodland,
ligh
was
slow
and
lu -
e ing,
much
mo e
like
ha
o
P.
au i us
(Fluckige
and
Beck,
1995).
Dis ance
T a elled
om
he
Roos
A
cons an
ea u e
o
he
beha iou
o
bo h
Pleco us
species
is
ha
hey
o age
close
o
oos s.
Ba a aud
(1990)
mis -ne ed
bo h
species,
and
only
e e
caugh
ba s
in
ne s
which
we e
se
up
close
o
human
dwellings.
Cas o
e
al.
(1993)
always
obse ed
P.
aus iacus
o -
aging
close
o
oos s,
and
Fluckige
and
Beck
(1995)
epo ed
ha
his
species
mo ed
a
maximum
o
1.4
km
om
hei
nu se y
oos
du ing
adio
acking.
Swi
and
Racey
(1983)
sigh ed
ma ked
P.
au i us
a
maximum
o
1.1
km
om
a
nu se y
oos
du ing
July
and
Augus
( he
lac a ion
pe iod)
and
Fuh mann
and
Sei z
(1992)
ound
adio- acked
ba s
o
his
species
mo ed
a
compa able
1.5
km
om
he
oos
du ing
lac a-
ion,
al hough
hey
o aged
up
o
3.3
km
om
he
oos in
Sep embe
and
Oc obe .
This
may,
howe e ,
ha e
been
due
o
a
sho age
o
a ailable
ood
close
o
he
oos
la e
in
sum-
me ,
since
En wis le
e
al.
(1996)
ound
no
educ ion
in
he
o aging
dis ance
o
emales
due
o
lac a ion.
En wis le
e
alls
ex ensi e
s udy
ound
ha
he
u hes
eeding
si e
was
2.8
km
om
he
main
oos
si e.
This
si e
was
used
by
a
male;
he
u hes
si e
used
by
a
emale
was
2.2
km
om
he
oos .
O e all,
ba s
spen
92%
o
hei
o aging
ime
wi hin
1.5
km
o
he
oos
(Figu e
4.5)
and
signi ican ly
mo e
o
hei
ime
wi hin
0.5
km
o
he
oos
han
u he
away.
When
da a
o
males
and
emales
we e
analysed
sepa a ely,
i
was
shown
ha ,
while
emales
spen
mos
o
hei
ime
wi hin
0.5
km
o
home,
males
showed
a
mo e
equal
dis-
ibu ion
o
hei
ime
be ween
dis ance
bands.
This
is
ob ious
in
Figu e
4.6,
which
shows
da a
o
pai s
o
male
and
emale
ba s
acked
simul aneously.
A
signi ican
posi i e
ela-
ionship
was
ound
be ween
he ime
spen
wi hin
0.5
km
o
he
oos
by
emale
P.
au i us
and
he
a ea
o deciduous
woodland
wi hin
ha
dis ance.
Thus,
i
appea s
ha
i
is
impo -
an
o
he
ba s
o
ha e
woodland
a ailable
close
o
he
oos ,
a
conclusion
which
ein o ces
he
inding
o
En wis le
e
al.
(1997)
ha
P.
au i us
selec
oos s
which
ha e
deciduous
woodland
wi hin
0.5
km,
a he
han
choosing
buildings
a
andom.
Fo aging
65
<0.5
0.5-1.0
1.0-1.5
>1.5
Dis ance
om
oos
(km)
FIG
4.5
The
pe cen age
o o al lying
ime
spen
by
16
adio- acked
P.
au i us
om
oos s
in
no h-eas
Sco land
in
dis ance
bands
p og essi ely
u he
om
oos s,
n
=
65
ba
nigh s
plus
7
hal
nigh s
( ep oduced
wi h
pe mission
om
En wis le
e
al.,
19%).
The
easons
why
emale
P.
au i us
o age
close
o
oos s
han
males
a e
no
clea .
I
is
possible
ha
emales
a e
es ic ed
because
hey
ha e
o
make
ene ge ically
expensi e
ligh s
back
o
he
oos
o
suckle
hei
in an s.
Al hough
his
explana ion
is
a
a iance
wi h
he
inding
o
En wis le
e
al.
(1996)
ha
he e
was
no
di e ence
be ween
he
dis ances
a -
elled
by
lac a ing
and
non- ep oduc i e
emales,
i
is
possible
ha
pa e ns
in ol ing
o ag-
ing
close
o
he
oos
may
become
in insic
in
emales
and
may
p e ail
e en
in
yea s
when
hey
do
no
gi e
bi h.
Al e na i e
explana ions
sugges ed
by
hese
au ho s
a e
ha
ei he
males
may
be
agg essi ely
excluded
om
close-by
o aging
a eas
o
ha
hey
may
bene i
om
educed
in aspeci ic
compe i ion
i
hey
mo e
u he
om
he
oos .

66
Fo aging
Beha iou
<0.5
0.5-1.0
1.0-1.5
>1.5
Dis ance
band
(km)
FIG
4.6
The
pe cen age
o
o al
lying
ime
spen
by
male
and
emale
P.
au i us
o
simul aneously
acked
pai s
in
dis ance
bands
p og essi ely
u he
om
oos s.
Ligh
ba s,
emale
ba s;
da k
ba s,
male
ba s
( ep oduced
wi h
pe mission
om
En wis le
e
al.,
1996).
Feeding
Ra e
and
Du a ion
o
Fo aging
Insec i o ous
ba s
eed
apidly,
pa icula ly
in
he
i s
hou
o
wo
o
o aging,
when
hey
a e
e y
hung y and
insec
a ailabili y
is
gene ally
high.
Ko un
and
Zhuko a
(1994)
es i-
ma ed
ha
mos
species
consume
app oxima ely
25%
o
hei
own
body
weigh
in
insec s
pe
nigh .
This,
o
a
P.
au i us
weighing
on
a e age
a ound
8
g,
means
i
ea s
abou
2
g
o
insec s,
mos ly
in
he
i s
2
h
o
o aging.
Since
la ge
c ane lies
weigh
abou
50
mg,
small
bee les
abou
75
mg
and
noc uid
mo hs
an
a e age
o
a ound
100
mg,
his
means
ha
a
Fo aging
67
long-ea ed
ba
would
ha e
o
ea
somewhe e
a ound
40
c ane lies,
27
small
bee les
(such
as
lea
o
ba k
bee les)
o
20
noc uid
mo hs
in
a
nigh .
Rydell
(1993)
sugges ed
ha ,
a
high
la i udes,
insec i o ous
ba s
may
ac
Pn™a"
Y
as
ene gy
maximize s
a he
han
as
ime
minimize s,
which
would
mean
hey
would
o age
du ing
all
he
ime
a ailable
i
i
we e
ene ge ically
p o i able
o
do
so
I
hey
we e
ime
minimize s,
he
du a ion
o
o aging
would
e lec
he
changing
ene gy
demands
o
ep o-
duc ion,
i.e.
o aging
ime
would
inc ease
slowly
du ing
p egnancy
and
hen
apidly
as
lac-
a ion
p oceeded.
No
e idence
was
ound
by
En wis le
e al.
(1996)
o
indica e
ha
his
was
he
case
in
P.
au i us
-
lac a ing
and
non- ep oduc i e
emales
and
males
all
had
o aging
du a ions
which
did
no
di e
signi ican ly
om
each
o he .
The
o al
median
o aging
ime
o
emales
was
235.7
min
and
ha
o
males
was
271.9
min.
These
we e
compa ab
e
o
he
a e age
o aging
du a ion
o
emales
in
Ge many
(353.6
min;
Fuh mann
and
Sei z,
1992).
I
he e o e
appea s
ha ,
a
high
la i ude,
sho ness
o
da kness
p ecludes
P.
au i us
om
being
a
ime
minimize .
Ins ead,
i
is
an
ene gy
maximize ,
o aging
o
as
long
as
da kness
pe mi s.
The e
is
e idence
ha
his
species
makes
ex ensi e
use
o
daily
o po
du ing
lac a ion
(Speakman
and
Racey,
1987;
see
also
Chap e
5)
o
make
up
o
he
ene gy
sho all
caused
by
he
demands
o
ea ing
an
in an .
CHAPTER
5
Rep oduc ion
"O
ATS
a e
placen al
mammals,
and
hey
sha e
he
ep oduc i e
cha ac e is ics
o
his
-L-J
g oup.
Ma ing
and
e iliza ion
a e
ollowed
by
a
pe iod
o
ges a ion,
du ing
which
he
g owing
emb yo
is
nou ished
by
he
placen a.
A e
he
young
is
bo n,
i
is
ca ed
o
and
ed
on
milk
by
i s
mo he
un il
weaning.
Howe e ,
ba s'
ep oduc i e
cycles
a e
a
om
ypi-
cal
o
small
mammals
—
ins ead
o
being
sho -li ed
and
p oducing
many
o sp ing,
like
mice,
ba s
li e
long
li es
and
p oduce
ela i ely
ew,
la ge
o sp ing
in
which
hey
in es
a
high
deg ee
o
ma e nal
ca e.
Ba s
such
as
Pleco us
species
which
li e
in
empe a e
zones
ha e
seasonal
cycles
which
a e
adap ed
o
inco po a e
a
pe iod
o
hibe na ion
and
o
ensu e
ha
he
young
a e
bo n
and
weaned
a
imes
o
yea
which
maximize
hei
chances
o su -
i al.
To
his
end,
hey
show
a
numbe
o
highly
specialized
ep oduc i e
adap a ions.
SEASONAL
CYCLE
Like
all
ba s
li ing
in
empe a e
zones,
long-ea ed
ba s
unde go
only
one
ep oduc i e
cycle
pe
yea .
They
ha e
a
ela i ely
long
ges a ion
pe iod
and
he
young
mus
be
bo n
as
ea ly
as
possible
in
summe
o
ensu e
hey
ha e
a
su icien
pe iod
a e
weaning
in
which
o
accumula e
enough
a
o
su i e
hei
i s
win e .
In
addi ion,
emales
need
an
adequa e
supply
o
insec
ood
o
sus ain
p egnancy
and
lac a ion.
Summe s
in
empe a e
zones,
pa -
icula ly
a
high
la i ude,
a e
no
long
enough
o
inco po a e
ma ing,
p egnancy,
weaning
and
p e-hibe nal
a
accumula ion.
The
monoes ous
cycle
is
he e o e
ex ended
o e
win-
e ,
wi h
ma ing
beginning
in
au umn
and
o ula ion
and
p egnancy
delayed
un il
sp ing
(Figu e
5.1).
The
p ocess
o
spe ma ogenesis
begins
in
la e
sp ing,
a e
males
ha e
eme ged
om
hibe na ion, and
con inues
h ough
summe .
Spe ma ozoa
a e
p oduced
la e
in
sum-
me ,
a e
which
he
es es
eg ess
in
Sep embe
and
Oc obe .
Ma ing
begins
in
Oc obe
and
con inues
spo adically
h ough
win e
and,
in
P.
au i us
bu
p obably
no
in
P.
aus i-
acus
(S ebbings,
1970),
in
ea ly
sp ing.
Spe ma ozoa
a e
s o ed
in
he
u e i
o
emales
and
in
he
epididymides
o
males
o e
win e .
Females
o ula e
in
la e
Ap il
o
May,
a e
hibe -
na ion
ends,
and
he
o um
is
e ilized
by
spe m
s o ed
in
he
u e us.
P egnan
ba s
mo e
in o
nu se y
oos s
du ing
sp ing
and
pa u i ion
akes
place
he e,
usually
in
ea ly
o
mid
July.
Mo he s
and
young
emain
in
nu se y
oos s
du ing
lac a ion
in
July
and
Augus
and
may
s ay
on
he e
in o
au umn.
The e
is
some
e idence
ha
ma ing
may
occu
in
nu se y
oos s
in
Pleco us
colonies.
The
ep oduc i e
cycle
is
he e o e
long
o
a
small
mammal
—
spe ma ozoa
p oduced
68
Male
Rep oduc ion
69
July
FIG
5.1
Schema ic
ep esen a ion
o
he
annual
ep oduc i e
cycle
o
long-ea ed
ba s
(adap ed
om
Racey,
1974a).
one
summe
do
no
esul
in
li e
young
un il
he
nex .
Since
he
as
majo i y_o
PUcoM
emales
p oduce
a
single
in an ,
he
ep oduc i e
a e
is
hus
e y
low.
Because
o
he
eng
and
complexi y
o
he
cycle,
a
emale
which
does
no
become
p egnan
which
abo s
a
e us
o
whose
in an
dies,
loses
i s
chance
o
ep oduce
o
a
whole
yea .
A
numbe
o
adap a-
ions
in
he
ep oduc i e
cycle
ha e
e ol ed
which minimize
he
chances
o
such
ailu es
These
ensu e
ha
a
high
p opo ion
o
adul
emales,
es ima ed
a
70%
(En wis le,
o
97%
(Benzal,
1991),
do
p oduce
young
each
yea
and
ha
he
in an s
ha e
he
bes
pos-
sible
chance
o
su i al.
MALE
REPRODUCTION
Rep oduc i e
O gans
The
ana omy
o
male
long-ea ed
ba s
is
ypical
o
espe ilionid
ba s
excep
ha
he e
a e
wo
pai s
o
Cowpe 's
glands
(Figu e
5.2)
ins ead
o
one.
The
possession
o
wo
pai s
o
hese
glands
appea s
o
be
sha ed
only
by
Pleco us
and
Co yno hinus,
a
closely
ela ed
No h
70
Rep oduc ion
Ame ican
genus
(Pea son
e
al.,
1952).
The
es es,
which
p oduce
spe ma ozoa,
consis
o
igh ly
coiled
semini e ous
ubules
su ounded
by
in e s i ial
cells
which
p oduce
he
ho -
mones esponsible
o
agg ession
and
libido.
The
es es
a e
abdominal
be o e
bi h
bu
descend
a ound
he
ime
o
bi h
o
occupy
a
posi ion
in
he
uppe
high,
le el
wi h
he
penis.
The
duc
sys em
includes
he
asa
e e en ia
and
epididymis,
which emp ies
in o
he
as
de e ens
and hen
he
u e h a.
The
accesso y
glands
include
he
ampulla y,
p os a e
and
Cowpe 's
glands.
All
o
hese
con ibu e
cons i uen s
o
he
semen,
including
essen ial
nu ien s,
uc ose
and
a ious
elec oly es
which
aid
in
main aining
he
pH
o
he
seminal
luid
a
a ound
7
(Hill
and
Smi h,
1984).
The
size
and
ac i i y
le els
o
he
accesso y
glands
a e
con olled
by
ho mones
sec e ed
by
he
in e s i ial
cells
in
he
es es
and
a y
h ough-
ou
he
annual
cycle.
The
penis
is
he
copula o y
o gan
and
i s
e ec ion
is
con olled
by
engo gemen
o
he
co po a
ca e nosa
wi h
blood.
Pleco us
also
ha e
a
baculum,
a
small
bone
which
caps
he
end
o
he
co po a
ca e nosa.
Ho monal
Con ol
o
he
Rep oduc i e
Cycle
Tes os e one,
sec e ed
by
he
es es,
con ols
he
p ocess
o
spe ma ogenesis
and
main ains
he
sec e o y
ac i i y
o
he
accesso y
glands.
The
ep oduc i e
cycle
o
male
long-ea ed
ba s
Male
Rep oduc ion
71
and
o
all
o
he
ba
species
which
hibe na e
a e
cha ac e ized
by
an
asynch onous,
annual
ec udescence
o
he
es es
and
accesso y
o gans,
and
his
asynch ony
esul s
in
cycles
in
bo h
which
a e
ou
o
phase
wi h
each
o he
(Gus a son
and
Damassa,
1987).
Al hough
he
o al
plasma
es os e one
le el
inc eases
a
he
same
ime
as
es icula
ac i i y
begins
in
sp ing,
he
accesso y
o gans
do
no
begin
o
inc ease
in
mass
o
show
o he
signs
o
seasonal
ac i i y
un il
some
ime
la e ,
when
plasma
es os e one
le els
a e
a
a
maximum.
This
delayed
esponse
by
he
accesso y
o gans
sugges s
ha
he e
is
some
o m
o
and ogen
esis-
ance,
which
appea s
o
be
due
o
a ia ions
in
and ogen
a ailabili y
a he
han
o
changes
in
he
sensi i i y
o
he
a ge
o gans.
Gus a son
and
Damassa
(1987)
sugges ed
ha
he
le -
els
o
plasma
es os e one
a e
con olled
by
a
sex
s e oid-binding
ho mone
(SBP)
which
has
been
ound
in
he
blood
o
se e al
espe ilionid
species
and
which
binds
plasma
es os-
e one,
ende ing
i
e ec i ely
una ailable
o
physiological
ac ion.
They
p oposed
ha
only
when
non
SBP-bound
le els
o
es os e one
inc ease
la e
in
summe
does
enough
ee
es os e one
ci cula e
o
cause
he
accesso y
o gans
o
become
ac i e.
Age
o
Ma u i y
S ebbings
(1966),
in
a
popula ion
s udy
o
Pleco us,
ound
ha
wo
ou
o
h ee
males
bo n
one
summe
unde wen
spe ma ogenesis
he
ollowing
summe
and
we e
hus
sexually
ma u e
(de ined
by
he
p oduc ion
o
spe m)
in
hei
second
au umn.
A
hi d
showed
no
signs
o
ma u i y
a
his
age
and
p esumably
did
no
unde go
spe ma ogenesis
un il
i
was
2
yea s
old.
Speakman
and
Racey
(1986)
epo ed
ha
3
ou
o
57
P.
au i us
males
cap u ed
in
no h-eas
Sco land
(i.e.
abou
5%)
showed
signs
o
sexual
ma u i y
in
he
au umn
o
hei
bi h,
when
hey
we e
abou
3
mon hs
old,
and
En wis le
e
al.
(in
p ess)
also
ound
ha
a
ew
showed
limi ed
es icula
de elopmen
a
3
mon hs,
al hough
mos
unde wen
spe ma ogenesis
o
he
i s
ime
in
he
summe
o
he
yea
ollowing
hei
bi h.
En wis le
e
al.
(in
p ess)
poin ed
ou
ha
he
deg ee
o
es icula
de elopmen
in
he
3
mon h
old
males
in
hei
s udy
was
much
less
han
ha
in
adul s
caugh
a
he
same
ime,
and
ha
hey
may
no
ha e
been
e ile.
Sexual
ma u i y
in
an
indi idual's
second
summe
hus
appea s
o
be
no mal
in
his
species,
as
in
mos
o he
espe ilionids.
A
simila
si ua ion
exis s
in
C.
ownsendii,
in
which
a
ew
males
ma u e
in
hei
i s
au umn
and
mos
in
hei
second
summe
o
au umn
(Pea son
e
al.,
1952),
and
in
Co yno hinus
a inesquii,
in
which
males
ma u e
in
hei
second
au umn
(Jones
and
Su kus,
1975).
I
seems
ha
a
ew
P.
au i us
may
no
ma u e
un il
a
leas
hei
hi d
yea
(S ebbings,
1966),
an
unusual
si ua ion
among
espe ilionid
ba s.
Fac o s
in luencing
a ainmen
o
sexual
ma u i y
Speakman
and
Racey
(1986)
ound
he e
was
a
s ong
di ec
ela ionship
be ween
he
age
a
which
male
P.
au i us
become
ma u e
and
hei
body
condi ion,
measu ed
by
means
o
an
index
(body
weigh / o ea m
leng h).
They
sugges ed
ha
long-ea ed
ba s
had
o
a ain
a
a ge
weigh
be o e
hey
a ained
pube y
and
ha
only
hose
in
e y
good
body
condi ion
would
ma u e
in
hei
i s
summe .
En wis le
e
al.
(in
p ess)
simila ly
ound
ha
ba s
which
showed
signs
o
es icula
g ow h
and
epididymal
de elopmen
a
3
mon hs
had
sig-
ni ican ly
highe
ela i e
body
condi ion
han
did
hose
which
emained
imma u e.
Benzal
(1991),
in
a
s udy
in
cen al
Spain,
ound
ha
ju enile
male
P.
au i us
gained
less
weigh
han
did
ju enile
emales
in
he
pe iod
be o e
hei
i s
hibe na ion.
He
sugges ed
ha
72
Rep oduc ion
mos
males
we e
no
hea y
enough
o
a ain
sexual
ma u i y
be o e
hey
wen
in o
hibe -
na ion.
The
ac o s
in luencing
age
o
ma u i y
a e
hus
likely
o
be
ime
o
bi h
(la e
bi hs
in
a
colony
will
esul
in
e y
ew
becoming ma u e
be o e
win e ),
wea he
condi ions,
ood
a ailabili y
and
o aging
abili y
in
indi idual
ba s.
Me hods
o
assessing
ma u i y
Because
he
es es
eg ess
seasonally
in
empe a e
zone
ba
species,
i
is
no
always
ob ious
which
males
o
adul
size
a e
ac ually
sexually
ma u e.
Racey
(1974b)
showed
ha
in
Pipis elluspipis ellus
assessmen
could
be
made
by
ex e nal
examina ion
ol
he
caudae
epi-
didymides.
The
unica
aginalis,
a
shea h
o
pe i oneum
co e ing
he
es es
and
epi-
didymides,
appea s
black
h ough
he
skin
due
o
he
p esence
o
pigmen -bea ing
cells,
o
melanocy es.
As
he
caudae
epididymides
become
dis ended
wi h
spe m,
he
shea h
s e ches
and
he
melanocy es
sepa a e
so
ha
he
colou
becomes
mo led,
and
he
unica
aginalis
emains
mo led
a e
he
spe m
ha e
been
oided;
i
does
no
egain
i s
je
black
colou
in
adul s.
Al hough
he
me hod
has
been
used
o de e mine
sexual
ma u i y
in
P.
au i us
(Speakman
and
Racey,
1986;
Boyd
and
S ebbings,
1989)
and
o he
species,
un il
ecen ly
i s
e icacy
has
no
been
es ed.
Howe e ,
En wis le
e
al.
(in
p ess)
ha e
now
epo ed
ha
26%
o
P.
au i us
males
we e
w ongly
assessed
by
he
me hod;
in
hei
s udy,
a
ew
males
known
o
be
ju eniles
had
comple ely
whi e,
unpigmen ed
caudae
epi-
didymides.
They
ound
ha
he
ela i e
size
and
shape
o
he
caudae
epididymides
was
a
mo e
eliable
indica o
in
P.
au i us
-
hose
o
imma u e
animals
we e
small
and
nodula ,
while
hose
o
adul s
we e
la ge
and
do so- en ally
la ened.
Spe ma ogenesis
The
p ocess
is
ini ia ed
in
la e
sp ing
o
ea ly
summe
in
mos
Eu opean
species,
and
con-
inues
h ough
summe .
A
he
end
o
summe ,
spe ma ozoa
a e
eleased
om
he
es es
in o
he
epididymides
and
he
es es
eg ess.
Tes icula g ow h
was
i s
obse ed
in
P.
au i-
us
in
Sco land
in
ea ly
July
(En wis le
e
al.,
in
p ess)
and
eached
a
peak
in
mid
Augus .
Changes
in
es icula
size
indica ed
ha
spe ma ogenesis
was
comple e
and
spe ma ozoa
we e
eleased
in o
he
epididymides
a ound
he
end
o
Augus .
The
es es
hen
eg essed,
bu
he
caudae epididymides
emained
la ge
o
some
mon hs,
be o e
g adually
sh inking
du ing
he
hibe na ion
pe iod.
The
es es
emained
quiescen
h oughou
hibe na ion.
The
ac o s
con olling
he
onse
o
spe ma ogenesis
a e
no
ully
unde s ood.
Racey
(1978)
epo ed
ha
p ema u e
a ousal
o
P.
pipis ellus
om
hibe na ion,
associa ed
wi h
bo h
ood
a ailabili y
and
empe a u e,
was
shown
o
esul
in
p ema u e ini ia ion
o
spe -
ma ogenesis.
Ligh
also
had
some
e ec ,
since
he
p ocess
was
inhibi ed
in
pipis elles
kep
in
o al
da kness,
bu
he e
was
no
e idence
ha
inc easing
pho ope iod
was
necessa y
o
s imula e
es icula
de elopmen .
Speakman
and
Racey
(1986)
ound
ha
a
ew
ju enile
P.
au i us
showed
signs
o
sexual
ma u i y
in
hei
i s
au umn
and
poin ed
ou
ha
hey
did
his
a
a
ime
when
dayleng h
was
becoming
sho e .
En wis le
e
al.
(in
p ess)
assessed
he
use
o
o po
by
male
P.
au i us in
summe
by
eco ding
he
p opo ion
o
o pid
males
caugh
in
oos s
and
in es iga ed
he
ela ionship
be ween
he
use
o
o po
and
es os e one
le els.
They
ound
ha
he
use
o
o po
was
lowes
in
July
and
Augus ,
when
spe ma ogenesis
was
a
i s
peak,
and
highes
in
May
and
Sep embe .
Use
o
o po
was
almos
comple ely
suspended
a
he
ime
when
he
ini ial
dis-
ension
o
he
caudae
epididymides
was
eco ded,
and
his
was
also
accompanied
by
a
d op
Male
Rep oduc ion
73
in
body
mass
and
a
endency
o
male
ba s
o
inc ease
hei
use
o
wa m
nu se y
oos s.
They
concluded
ha
o po
and
peak
spe ma ogenesis
may
be
incompa ible
in
his
species,
ei he
because
endo he my
is
a
p e equisi e
o
spe ma ogenesis
o
because
ba s'
abili y
o
en e
o po
is
educed
when
es os e one
le el
is
e y
high.
The
wo
explana ions
a e
no
mu ually
exclusi e.
A e
spe ma ogenesis
is
comple e,
es os e one
le el
d ops
and
he
accesso y
o gans
a e
main ained
by
much
lowe
le els.
I
appea s
ha
his
lowe
le el
does
no
inhibi
hibe na ion,
since
a
mode a e
le el
is
main ained
all
win e .
En wis le
e
al.
(in
p ess)
sugges ed
ha
le els
o
es os e one
may
b ie ly
be
ele a ed
du ing
pe iodic
a ousals
om
hibe na ion
in o de
o
main ain
seconda y
ep oduc i e
p ocesses.
Spe m
S o age
by
Males
Spe m
s o ed
in
he
caudae
epididymides
o e
win e
a e
capable
o
e iliza ion
in
a
num-
be
o
species
o
hibe na ing
ba s.
S elko
(1962)
showed
ha
an
inc easing
p opo ion
o
emales
o
ou
espe ilionid
species,
including
P.
au i us,
we e
insemina ed
as
hibe na ion
p og essed.
Racey
(1972)
conduc ed
expe imen s
on
cap i e
noc ules
(Nyc alus
noc ula),
in
which
emales
we e
no
allowed
o
ma e
in
au umn,
bu
we e
in oduced
o
males
a
a i-
ous
imes
du ing
win e
and
hen
became
p egnan
he
ollowing
sp ing.
The
minimum
s o age
ime
by
a
male
was
aken
as
being
he
in e al
be ween
he
da e
o
dis ension
o
he
caudae
epididymides
and
he
da e
when
he
emale
was
in oduced.
In
his
way,
Racey
showed
ha
spe m
emained
capable
o
e iliza ion
o
a
leas
3
mon hs
and
up
o
7
mon hs.
A
his ological
s udy
o
ep oduc i e
ac s
aken
om
hibe na ing
ba s
(Racey,
1975)
e ealed
no
di e ences
be ween
noc ules
and
a
a ie y
o
o he
B i ish
species
includ-
ing
P.
au i us
in
he
s o age
si es
o
appea ance
o
s o ed
spe m,
so
i
is
likely
ha
he
ia-
bili y
o
spe m
o
all
hese
species
is
simila .
In
all
o
hem,
spe ma ozoa
we e
o ien a ed
owa ds
he
epi helial
cells
o
he
as
de e ens,
and
i
is
p obable
ha
hei
su i al
was
in
some
way
connec ed
wi h
hese
cells.
Ma ing
S elko
(1962)
caugh
emale
P.
au i us
in
hibe nacula
in
a i icial
ca es
du ing
win e .
He
ex ac ed
he
u e i,
washed
ou
he
con en s
and
examined
hem
mic oscopically
o
s o ed
spe ma ozoa.
In
No embe ,
sho ly
a e
hei
a i al
a
he
ca e,
only
14%
o
emales
we e
insemina ed.
In
Decembe ,
his
had
isen
o
63%,
in
Janua y-Ma ch
o
81%,
and
by
Ap il
100%
we e
insemina ed.
He
concluded
ha
in
his
species
ma ing
occu ed
h oughou
win e ,
ei he
h ough
males
a ousing
and
copula ing
wi h
do man
emales
o
du ing
pe i-
odic
a ousals
and
mo es
o
di e en
pa s
o
he
ca e
by
g oups
o
ba s.
Copula ion
by
males
wi h
o pid
emales
has
also
been
eco ded
in
C.
ownsendii
by
Pea son
e
al.
(1952).
S elko 's
da a
indica e
ha
he
main
ma ing
season
o
P.
au i us
is
win e
and
ea ly
sp ing,
a he
han
au umn
as
is
he
case
in
mos
espe ilionid
species.
S ebbings
(1970)
obse ed
ma ings
in
P.
au i us
in
Sep embe -Oc obe
and
in
Ap il
wi h
equal
equency,
bu
epo ed
ha
P.
aus iacus
om
he
same
oos
ma ed
only
in
au umn.
Ho acek
(1975)
simila ly
epo ed
di e ences
in
he
ma ing
beha iou
o
he
wo
species.
In
P.
au i us,
ma -
ing
mainly
occu ed
in
win e - ype
oos s
and
ac i i y
in
ansi o y
oos s
(Chap e
6)
was
high,
pa icula ly
in
sp ing,
while
in
P.
aus iacus
i
occu ed
mainly
in
summe
oos s
and
ansi o y
oos s
we e
no
used.
P.
aus iacus
hus
do
no
appea
o
ma e
in
sp ing, al hough
86
Rep oduc ion
condi ions.
The
wo
me hods
we e
employed
simul aneously
on
he
same
ba s,
and
calo ime y
was
assumed
o
gi e
he
'co ec '
measu emen .
They
ound
ha
DLW
consis-
en ly
o e es ima ed
ca bon
dioxide
p oduc ion
by
abou
14%
and
concluded
ha
he e
was
a
sys ema ic
bias
in
he
echnique.
This
was
o se
by
he
ac
ha
DLW
p o ided
in aluable
da a
which
could
no
be
collec ed
in
he
ield
in
any
o he
way.
In
addi ion,
o
he
pu pose
o
compa ing
ene gy
expendi u e
a
di e en
s ages
o
ep oduc ion,
bias
is
no
a
se ious
p oblem
p o ided
i
is
cons an .
S a egies
o
Accommoda ing
Ene gy
Demand
The e
a e
h ee
ways
in
which
animals
a e
known
o
espond
o
he
inc eased
ene gy
demand
o
p egnancy
and
lac a ion:
•
hey
may
u ilize
s o ed
a
o
ood
hoa ds
•
hey
may
inc ease
ood
in ake
•
hey
may
educe
expendi u e
on
some
componen
o
hei
ene gy
budge
no
con-
ce ned
wi h
ep oduc ion;
his
is
e e ed
o
as
compensa ion.
No
ba
species
is
known
o
hoa d
ood
and
s o age
as
a
is
mainly
con ined
o
la ge
mammals,
al hough
some
species
o
small
ones
do
deposi
a
which
is
used
in
lac a ion.
In
ba s,
a
accumula ion
has
he
p oblem
ha
ca ying
ex a
weigh
on
o aging
ligh s
in ol es
ex a
ene gy
expendi u e.
The e
is,
in
any
case,
a
limi
o
hei
ca ying
capaci y
and
hey
al eady
ha e
o
anspo
a
e us
o ,
on
occasions,
a
la ge
in an .
The e
is
e idence
(Speakman
and
Racey,
1987)
ha
P.
au i us
do
inc ease
hei
a
ese es
du ing
p egnancy,
bu
i
was
es ablished
ha
he
s o ed
a
could
p o ide,
a
mos ,
2-3
days'
wo h
o
ex a
ene gy.
Speakman
and
Racey
sugges ed
ha
a
deposi ion
in
his
species
is
mo e
likely
o
be
an
insu ance
agains
sudden,
se ious
sho alls
in
ene gy
supply
han
a
p olonged
subsidy
o
ep oduc i e
equi emen s.
Inc eased
ood
consump ion
o
compensa e
o
high
ene gy
demand
has
been
widely
demons a ed
in
small
e es ial
mammals,
bu
he e
a e
a
numbe
o
easons
why
i
is
less
likely
o
be
used
by
ba s.
The
ime
a ailable
o
o aging
is
limi ed
by
sho
nigh s,
especially
a
high
la i ude,
and
he e
a e
also
empo al
limi a ions
on
ood
a ailabili y
-
insec
num-
be s
all
a
ce ain
imes
du ing
he
nigh
(Chap e
3).
The
high
ene ge ic
cos
o
ba
ligh ,
es ima ed
a
21
imes
he
basal
me abolic
a e
o
P.
au i us
by
Racey
and
Speakman
(1987),
means
ha
ex ending
he
o aging
pe iod
would
only
be
an
e icien
me hod
o
assimila ing
ex a
ene gy
i
p ey
le els
we e
easonably
high.
Finally,
he
amoun
o
ood
a
ba
can
ea
is
limi ed
by
he
handling
ime,
which
is
he
ime
aken
o
ack
down,
ca ch
and
consume
a
p ey
i em.
Swi
(1980)
and
Maie
(1992)
ound
changes
in
he
ac i i y
pa -
e ns
o
P.
pipis ellus
in
p egnancy
and
lac a ion,
bu
no
inc ease
in
he
o al
ime
hese
ba s
spen
o aging
in
a
nigh .
In
P.
au i us,
En wis le
e
al.
(1996)
ound
ha
nei he
he
ime
spen
o aging
no
he
dis ance
indi iduals
mo ed
om
he
oos
di e ed
be ween
lac a -
ing
and
non- ep oduc i e
emales.
The e
was
hus
no
e idence
o
inc eased
ood
consump-
ion
du ing
lac a ion.
Because
insec i o ous
ba s
in
empe a e
egions
a e
he e o he mic,
hey
can
use
o po
o
educe
ene gy
demand
as
a
o m
o
compensa ion
-
an
op ion
open
o
ew
small
e es ial
mammals.
The e
is
e idence
om
a
numbe
o
s udies
ha
he
use
o
o po
is
widesp ead
among
insec i o ous
ba s.
Speakman
and
Racey
(1987)
used
he
DLW
echnique
o
Pla e
1
A
b own
long-ea ed
ba
[Pleco us
au i us)
in
ligh .
The
la ge,
e ec
ea s
can
be
seen
in
o chligh .
Pho o:
F ank
G eenaway.
m

Pla e
2
G ey
long-ea ed
ba
(Pleco us
aus iacus).
The
agus
shape,
da k
b own
acial
colou ing
and
sho
humbs
iden i y
his
indi idua
.
Pho o:
F ank
G eenaway.
Pla e
3
{below)
A
b own
long-ea ed
ba
a
oos
wi h
ea s
in
he
' am's
ho n'
posi ion.
The
ans e se
olds
in
he
conch
close
in
a
an-like
ac ion
as
he
ea
elaxes.
Pho o:
John
Haddow.
n-x x
-
- i
'
"
Pla e
4
Because
long-ea ed
ba s
ly
inside
oos s,
hei
d oppings
become
sca e ed
a ound
a ics
and
may
co e
u ni u e
o
o he
possessions
s o ed
he e.
Pla e
5
(below)
P.
au i us
lying
close
o
ege a ion.
Du ing
oliage
gleaning,
b own
long-ea ed
ba s
equen ly
s op
echoloca ing
and
sea ch
o
p ey
by
passi e
lis ening.
Pho o:
F ank
G eenaway.
Pla e
6
Ins
emains
.accumula e
nea h
eeding
pe ch
s.
Clea ly,
he
ba
ha
used
his
pe ch
had
been
hun ing
la ge
yellow
unde wing
mo hs.
Pho o:
F ank
G eenaway.
Pla e
7
Flyways
a e
consis en ly
used
by
long-ea ed
ba s
o
a oid
c ossing
open
spaces
while
commu ing
be ween
oos s
and
o aging
a eas.
They
ly
close
o
ege a ion
along
landscape
ea u es
such
as
o e g own
s eams.
Pla e
8
A
g oup
o
b own
long-ea ed
ba s
in
hei
no mal
oos ing
posi ion
in
an
a ic.
They
we e
pho og aphed
om
below
and
a e
huddled
be ween
he
idge
beam
and
he
sa king,
in
he
angle
o med
by
a
a e .
Pla e
10
(below)
B own
long-
ea ed
ba s
ha e
been
ound
o
selec
olde
houses
wi h
complex
oo
spaces.
A
colony
o
80
oos ed
in
his
Vic o ian
house
and
equen ly
mo ed
be ween
oo
compa men s.
Pla e
9
A
emale
b own
long-ea ed
ba
wi h
a
newbo n
in an
Babies
a e
con inuous!
a ached
o
hei
mo he 's
nipple
du ing
hei
i s
week
o
li e.
Pho o:
F ank
G eenaway.
Pla e
11
(abo e)
Long-ea ed
ba
oos s
a e
likely
o
be
si ua ed
in
wooded
alleys.
Well-main ained
ishing
i e s
a e
an
impo an
ea u e
o
he
ecology
o
he
species.
Pla e
12
(below)
P.
au i us
selec
houses
which
ha e
woodland
wi hin
0.5
km.
This
oos
is
su i
ounded
by
ma u e
deciduous
and
coni e ous
ees.
Pla e
13
A
hibe na ing
b own long-ea ed
ba
wi h
olded
ea s.
The
agi
hang
down,
supe icially
esembling
he
ea s
o
o he
species.
Pho o:
F ank
G eenaway.
Ene ge ics
o
Rep oduc ion
87
measu e
ene gy
expendi u e
in
ee-li ing
P.
au i us
h oughou
he
summe
in
no h-eas
Sco land.
Ba s
we e
cap u ed
in
nu se y
oos s,
weighed
and
measu ed
and
hei ep oduc-
i e
s a e
asce ained.
They
we e
hen
injec ed
pe i oneally
wi h
wa e
con aining
labelled
oxygen
and
hyd ogen.
A e
allowing
he
l8oxygen
90
min
o
each
equilib ium,
a
blood
sample
was
aken
om
each
ba
and
hey
we e
hen
eleased
in
he
oos .
Twen y- ou
hou s
la e
hey
we e
ecap u ed,
eweighed
and
a
second
se ies
o
blood
samples
ob ained.
The
s udy
ound
ha
he
body
mass
o
emales
was
highe
(mean=9.2
g)
a
he
s a
o
lac-
a ion
han
a
he
s a
o
p egnancy
(mean=7.3
g),
indica ing
ha
a
signi ican
inc ease
had
occu ed
du ing
p egnancy
and
ha
his
was
no
a ibu able
o
he
mass
o
he
e us
and
accesso y
s uc u es.
I
was
pa ly
due
o
enla gemen
o
he
mamma y
glands,
bu
his
explained
only
10%
o
he
inc ease.
The
au ho s
sugges ed
ha
he
es
may
ha e
been
a ,
o
be
used
as
an
eme gency
supply
which
could
ide
he
ba
o e
in
he
e en
o
a
2-3
day
se ious
ood
sho age.
Speakman
and
Racey
nex
p edic ed
he
ene ge ic
cos s
in
p egnancy
and
lac a ion
o
a
ba
which
emained
con inuously
endo he mic,
p edic ions
being
made
om
esul s
o
labo a o y
espi ome y
combined
wi h
mic oclima e
measu emen s
in
he
ield
and
allome ic
p edic ions
o
ligh
cos .
These
we e
compa ed
wi h
he
ield
esul s
om
he
s udy
(Figu e
5.6).
In
ea ly
p egnancy,
he
obse ed
expendi u e
was
abou
50%
o
he
p edic ed
alue
o
an
endo he mic
ba ,
sugges ing
ha
daily
o po
was
being
used.
The
obse ed
ene gy
expendi u e
hen
inc eased
apidly,
un il
by
pa u i ion
i
exceeded
he
p edic ed
a e
o
an
endo he mic
ba .
This
sugges ed
ha
ba s
shi ed
s a egy
du ing
la e
p egnancy
o
become
con inuously
endo he mic.
Immedia ely
a e
gi ing
bi h,
ene gy
expendi u e
d opped
again
o
a
much
lowe
han
expec ed
le el,
sugges ing
ha
hey
compensa ed
o
high
ene gy
demand
by
ex ensi e
use
o
o po .
The
s a egy
used
by
long-
ea ed
ba s
is
hus
complex.
The
con inuously
endo he mic
pe iod
in
la e
p egnancy
sug-
ges s
he
possibili y
ha
o po
a
his
s age
may
be
incompa ible
wi h
e al
de elopmen
(Speakman
and
Racey,
1987),
o
possibly
he
delay
in
ges a ion
which
would
esul
om
o po
would
mean
ha
he young
would
be
bo n
oo
la e
in
he
yea
o
ha e
a
ealis ic
chance
o
accumula ing
enough
a
o
su i e
hei
i s
win e .
The e
does
no
appea
o
be
a
compa able
pe iod
o
endo he my
du ing
lac a ion,
so
milk
p oduc ion
mus
be
compa -
ible
wi h
daily
o po .
E ec
o
To po
on
Milk
P oduc ion
The
disad an age
o
a
emale
ba
o
sa ing
ene gy
by
he e o he my
is
he
non-speci ici y
o
o po .
Mamma y
me abolism
is
slowed
down,
as
is
ha
o
o he
body
issues
and
he
esul
would
be
a
dec ease
in
milk
sec e ion
unless
he
gland
we e
p o ec ed
in
some
way.
Wilde
e
al.
(1995)
in es iga ed
lac a ion
in
P.
pipis ellus
and
ound
ha
milk
p oduc ion
was
ai-
lo ed
o
he
in e mi en
suckling
pa e n
o
he
species
by
local
mechanisms
wi hin
he
mamma y gland,
which
esponded
o
he
equency
and
comple eness
o
milk
emo al.
Reduced
body
hea
did
dec ease
he
a e
o
milk
syn hesis
and
he e o e
milk
p oduc ion
was
educed
while
ba s
we e
o pid.
Howe e ,
he
esponse
o
he
mamma y
gland
o
e-
quen
o
in e mi en
suckling
was
an
acu e
one,
unlike
ha
in
mos
o he
mammals,
in
which
i
is
a
long- e m
esponse.
This
means
ha ,
in
pipis elles,
milk
syn hesis
is
inc eased
by
equen
e u ns
o
he
oos
o
suckle
in an s.
I
a
simila
si ua ion
exis s
in
Pleco us
species,
hen
his
would
explain
why
emales
e u n
o
hei
nu se y
oos s
se e al
imes
pe
nigh
(En wisde
e al.,
1996;
see
Chap e
4)
du ing
he
lac a ion
pe iod.
A e
each
isi ,
he
88
Rep oduc ion
10
ea ly
mid
P egnancy
la e
ea ly
mid
Lac a ion
la e
FIG
5-6
Va ia ions
in
daily
ene gy
expendi u e
in
ee-li ing
emale
P.
au i us
du -
ing
p egnancy
and
lac a ion,
compa ed
wi h
p edic ed
alues
o
a
con inuously
endo he mic
ba
(see
ex ).
Measu emen s
we e
made
using
he
DLW
echnique,
and
each
poin
ep esen s
a
single
indi idual
( ep oduced
wi h
pe mission
om
Speakman
and
Racey,
1987).
ee-li ing
ba s
measu ed
by
DLW;
A-A,
p edic ed
alues
o
a
con inuously
endo he mic
ba .
acu e
esponse
o
he
mamma y
gland
o
suckling
p obably
causes
apid
milk
p oduc ion
du ing
he
nex
bou
o
o aging.
In
his
way,
an
in an could
ecei e
mos
o
i s
daily
sup-
ply
o
milk
in
wo
o
h ee
eeds
and
milk
p oduc ion
could
hen
be
g ea ly
educed
while
he
mo he
is
o pid.
CHAPTER
6
Social
O ganiza ion
and
Beha iou
R
OOSTS
a e
an
impo an
aspec
o
he
ecology
o
long-ea ed
ba s.
They
p o ide
shel-
e
om
he
wea he ,
p o ec ion
om
p eda o s
and
a
wa m
en i onmen
in
which
o
p oduce
and
ea
young.
How
oos s
a e
selec ed,
how
ba s
use
hem
and
whe he
and
why
hey
mo e
be ween
oos s
in
an
a ea
a e
all
ques ions
which
ha e
been
in es iga ed
ecen ly.
This
chap e
also
looks
a
he
s uc u e
o
nu se y
colonies,
a
whe e
males
li e
du ing
sum-
me
and
how
his
a ec s
ma ing
beha iou
and
a
beha iou al
in e ac ions
be ween
mo he s
and
hei
o sp ing.
ROOSTING
BEHAVIOUR
Roos
Si es
In
common
wi h
mos
ba
species,
Pleco us
do
no
exca a e
o
build
places
o
li e,
bu
ely
o
shel e
on
exis ing
s uc u es.
In
p ehis o ic
imes
hese
s uc u es
we e
na u al
ones
such
as
ca es,
ee
holes
and
ock
c e ices,
bu
sinan h opic
(a i icial)
oos s
ha e
become
e y
impo an
o
many
ba s.
Some
species,
such
as
P.
pipis ellus,
ha e
become
so
well
adap ed
o
li ing
in
sinan h opic
oos s
ha
hey
a e
a ely
ound
anywhe e
else.
P.
au i us
in
B i ain
also
seem
o
ely
hea ily
on
such
oos s,
since
hey
a e
one
o
he
B i ish
species
mos
closely
associa ed
wi h
a ics.
Sinan h opic
oos s
ha e
eplaced
many
na u al
s uc-
u es;
hey
include
oo
spaces
eplacing
ee
ca i ies;
c acks
in
buildings,
b idges
and
expansion
join s
eplacing
ock
c e ices;
and
mines,
unnels
and
cul e s
eplacing
ca es.
P.
au i us
Roos s
The
na u al
oos s
o
his
species
a e
hough
o
be
ee
holes
(Ho acek,
1975),
and
in
cen-
al
Eu ope
he e
is
e idence
ha
hey
a e
s ill
used
ex ensi ely
as
summe
oos s
(Baue ,
1960;
Hanak, 1969).
P.
au i us
is
one
o
he
Eu opean
species
mos
likely
o
use
ba
boxes
(Chap e
9),
pa icula ly
in
coni e ous
o es s
whe e
he e
a e
ew
buildings
and
na u al
ee
holes
a e
sca ce.
Ba
boxes
esemble
ee
holes
and
so
hey
a e
used
as
subs i u es.
In
Swi ze land,
P.
au i us
a e
mo e
o en
ound
in
ee
holes
han
in
houses
(Fluckige
and
Beck,
1995),
and
S u z
(1989)
epo ed
ha
hey
ended
also
o
oos
a
high
al i ude
away
om
cen es
o
human
popula ion.
Howe e ,
in
o he
a eas
(e.g.
cen al
Bohemia;
Ho acek,
1975),
b own
long-ea ed
ba s
o m
nu se y
colonies
almos en i ely
in
houses,
despi e
he
ac
ha
he e
a e
plen y
o
a ailable
ee
holes.
Ha ma a
(1973)
also
ound
hem
mainly
in
89

90
Social
O ganiza ion
and
Beha iou
old
buildings
in
he
Czech
Republic.
In
Sco land,
En wis le
(1994)
ound
ha
nu se y
oos s
we e
almos
all
in
houses
and
ha
ee holes,
oge he
wi h
a m
buildings,
we e
used
only
as
empo a y
oos s
a
imes
when
ood
was
in
sho
supply
and
ba s
became
o pid
o
sa e
ene gy.
I
is
possible
ha
sinan h opic
oos s a e
used
mo e
by
his
species
a
high
la -
i ude
because
he
wa m h
hey
p o ide
is
c i ical
o
he
ba s
abili y
o
p oduce
and
eai
young
in
cool,
sho
summe s.
A ics
also
ha e
an
ad an age
o e
ee
holes
in
ha
hey
p o-
ide
space
o
young
ba s
o
p ac ise
lying
be o e
hey
ha e
o
go
ou side
o
he
i s
ime.
Chu ch
lo s
ha e
been
epo ed
o
be
used
as
oos
si es
in
he
Ne he lands
(Daan,
1980)
and
P.
au i us
also
make
use
o
chu ches
and
chu ch
po ches
in
England.
One
such
po ch
was
used
as
a
day
oos
o
a
colony
and
ano he
only
as
a
nigh
oos
(B own
e
al.,
1983).
Such
oos
si es
a e
usually
in
he
po ch
oo ,
agains
he
main
wall
o
he
chu ch
and
sou h- acing.
P.
aus iacus
Roos s
Unlike
hei
congene ics,
g ey
long-ea ed
ba s
almos
always
use
sinan h opic
oos s
du ing
summe
and
gene ally
li e
in
close
p oximi y
o
human
se lemen s
han
do
P.
au i us
(Hanak,
1969).
The
use
o
ee
holes
by
nu se y
colonies
o
his
species
is
unknown
and
ba s
li e
only
in
a ics
(Ho acek,
1973;
Gaisle
e
al.,
1990;
Fluckige
and
Beck,
1993).
These
a e
always
wa m
and
in
old
buildings
(Ha ma a,
1973);
colonies
ha e
been
epo ed
o
use
chu ch
lo s
which
hey
equen ly
sha e
wi h
colonies
o
P.
au i us
(Daan,
1980).
These
indings
suppo
he
heo y
o
Ho acek
(1975)
ha
P.
aus iacus
is
a
ecen
inhabi-
an
o
Eu ope
which
sp ead
ac oss
he
con inen
only
in
his o ical
imes
as
wa m,
sinan-
h opic
oos s
became
a ailable.
Posi ion
o
Ba s
wi hin
Roos s
B own
and
g ey
long-ea ed
ba s
di e
in
he
posi ions
in
which
hey
a e
mos
commonly
ound
in
oos s
(Ho acek,
1975).
P.
au i us
was
desc ibed
by
Gaisle
(1966)
as
a
issu e
oos e
which
usually
main ains
con ac
wi h
oo
beams
on
wo
sides.
The
commones
place
o
ind
hem
is
in
he
apex
o
he
oo ,
in
he
angle
be ween
he
idge
beam
and
he
a e s
( he
diagonal
beams
which
p ojec
downwa ds
om
ei he
side
o
he
idge
owa ds
he
sides
o
he
a ic).
They
a e
also
some imes
ound
a
gable
ends
be ween
s one
walls
and
beams
(Figu e
6.1).
P.
aus iacus,
on
he
o he
hand,
has
been
desc ibed
as
a
space
oos ing
species
(Gaisle ,
1966)
which
oos s
in
con ac
wi h
wood
on
one
side
only.
Ho acek
(1975)
main ained
ha
hey
a e
mos
equen ly
ound
in
he
apex
o
he
oo ,
on
he
idge
beam
be ween
join s
wi h
a e s
and
no
in
hei
angles
(Figu e
6.1).
Less
equen ly
hey
a e
ound
in
he
angles
o
he
a e s.
This
di e ence
suppo s
he
heo y
(Ho acek,
1975)
ha
he
wo
species
had
di e en
o igins
and
di e en
p e-sinan h opic
oos s:
P.
au i us
used
ee
holes,
bu
P.
aus iacus
p obably
oos ed
in
la ge
ca es.
Roos
Selec ion
Long-ea ed
ba s
ely
hea ily
on
hei
oos s
o
p o ide
bo h
wa m h and
p o ec ion
and
hus
he
cha ac e is ics
o
he
oos
may
exe
an
impo an
in luence
on
he
su i al
o
a
92
Social
O ganiza ion
and
Beha iou
colony.
Some
buildings
may
p o ide
mo e
sui able
condi ions
han
o he s
and
i
seems
likely
ha
colonies
selec
he
buildings
in
which
hey
oos .
En wis le
e al.
(1997)
se
up
a
s udy
o
in es iga e
his.
They
loca ed
a
o al
o
54
P.
au i us
oos s
in
hei
s udy
a ea
in
no h-eas
Sco land
and
hen
compa ed
cha ac e is ics
o
hese
oos s
wi h
hose
o
o he
buildings
andomly
selec ed
om
a
elephone
di ec o y
o
he
a ea.
The
s udy
ound
ha
he
use
o
houses
as
oos
si es
was
no
andom;
oos
selec ion
was
occu ing
and
ba s
appea ed
o
selec
houses
bo h
o
hei
loca ion
and
su ounding
habi a
and
o
speci ic
physical
ea u es
o
he
building.
Habi a
Mo e
oos s
han
expec ed
we e
si ua ed
in
i e
alleys
and,
compa ed
wi h
he
andomly
picked
buildings,
oos s
we e
signi ican ly
close
o
woodland
and
o
open
wa e .
The
median
dis ance
o
he
nea es
woodland
was
60
m
(«=48)
o
oos s
and
215
m
{n—16)
o
andom
houses.
Fu he mo e,
he e
we e
signi ican ly
g ea e
a eas
o
bo h
deciduous
and
coni e ous
woodland
wi hin
0.5
km
o
oos s
han wi hin
his
dis ance
o
andom
houses.
Woodland
close
o
oos s
is
e y
impo an
(Chap e
4),
bo h
because
P.
au i us
spend
mos
o
hei
o aging
ime
in
woodland
less
han
0.5
km
om
he
oos
and
because
such
close-by
woodland
p o ides
p o ec ion
om
p eda o s
and
allows
ba s
o
eme ge
o
o age
ela i ely
ea ly
in
he
e ening.
Roos s
may
ha e
been
mainly
in
i e
alleys
because,
in
Sco land,
his
is
whe e
mos
woodland
is
concen a ed.
Well-main ained
salmon
ishing
i e s
(such
as
he
Dee
and
he
Spey,
bo h
o
which
a e
in
he
s udy
a ea)
usually
ha e
ees
along
hei
banks,
wi h
especially
high
concen a ions
o
deciduous
woodland.
In
con as ,
much
o
he
coun yside
away
om
i e s
is
pas u e
o
moo land
wi h
ew
ees.
Simila ly,
he
inding
ha
oos s
we e
close
o
open
wa e
han
we e
andomly
picked
houses
may
ha e
been
due
o
he
si ua ion
o
oos s
in
wooded
i e
alleys.
The
median
dis ance
om
oos s
o
he
nea es
open
wa e
was
215
m
(«=48)
and
ha
om
andom
houses
was
450
m(«=16).
Roos s
we e
ound
o
be
associa ed
wi h
illages,
p obably
e lec ing
he
dis ibu ion
o
buildings
—
a eas
con aining
mo e
houses
p esen ed
mo e
oos ing
oppo uni ies.
Howe e ,
no
oos s
we e
loca ed
wi hin
Abe deen
ci y,
p obably
because
he
lack
o
woodland
in
a
ci y
would
make
i
unsui able
o
b own
long-ea ed
ba s.
Hanak
(1969),
Gaisle
(1979)
and
Gaisle
and
Baue o a
(1985-6)
all
simila ly
epo ed
ha
P.
au i us a oided
u ban
a eas.
Physical
ea u es
o
he
building
Compa ed
wi h
adjacen
houses
no
used
as
oos s,
En wis le
e
al.
(1997)
epo ed
ha
hose
used
as
oos s
we e
olde
and
con ained
la ge
oo
spaces
which
we e
mo e
likely
o
be
di ided
in o
a
numbe
o
compa men s.
Se en y- h ee
pe cen
o
a ics
used
as
oos s
had
mo e
han
one
compa men
( ange
=
1-8),
which
ga e
ba s
a
ange
o
oos
si es,
and
hus
a
ange
o
oos
empe a u es
om
which
o
selec
a
si e.
Roos s
we e
also
mo e
likely
o
be
lined
wi h
ough
wood
and
o
con ain
sa king
(boa ds
which
co e
he
a e s
and
on
which
he
oo
sla es
es ).
They
we e
less
likely
o
con ain
glass
ib e
insula ion
ma e ial
han
we e
adjacen
houses.
These
ea u es
we e
p obably
conce ned
wi h
wa m h,
since
a ics
wi h
sa king
and
wi hou
insula ion
will
main ain
a
ela i ely
high
empe a u e,
al hough
i
is
possible
ha
ough
wood
lining
was
also
chosen
by
he
ba s
because
i
esem-
bled
hei
na u al
oos
si es
in
ees.
La ge
oo
spaces
may
ha e
been
chosen
because
P.
au i us
need
enough
space
o
ly
in
he
oos ,
al hough
his
ea u e
was
no
signi ican
when
The mo egula ion
in
Roos s
93
con olled
o
by
loca ion
o
he
oos
(En wis le,
1994).
Wooden
lining
and
la ge
oo
spaces
wi h
se e al
compa men s
a e
gene ally
ea u es
o
olde
houses,
which
explains
he
s udy's
inding
ha
P.
au i us
selec ed
houses
ha
we e,
on
a e age,
abou
100
yea s
old
(En wis le,
1994).
This
con as s
wi h
houses
in
which
pipis elles
oos ,
which
a e
an
a e -
age
o
15.4
yea s
old
(Wa dhaugh,
1992).
Pipis elles
ha e
di e en
oos ing
equi emen s,
which
a e
ul illed
by
mode n
house
design.
P.
au i us
oos
in
he
da kes
a ailable
pa s
o
a ics
(En wis le,
1994),
al hough
hey
do
ole a e
some
deg ee
o
illumina ion,
no mally
om
small
skyligh
windows.
Such
low
le el
ligh
may
e en
be
bene icial
o
hem
by
allowing
hem
o
moni o
when
i
is
da k
enough
o
go
ou
and
o age.
The
absence
o
cobwebs
gene ally
associa ed
wi h
Pleco us
oos s
(Chap e
2)
is
p obably
due
o
hem
being
emo ed
ei he
by
ai
cu en s
c ea ed
by
he
ba s'
ligh
o
by
physical
con ac
wi h
hei
wings,
since
En wis le
(1994)
ound
no
e i-
dence
ha
a ics
wi h
many
cobwebs
we e
a oided.
The e
was
also
no
e idence
ha
ba s
dese ed
oos s
as
he
olume
o
hei
d oppings
inc eased,
al hough
En wis le
(1994)
poin ed
ou
ha ,
since
P.
au i us
gene ally
o m
small
colonies,
he
olume
o
guano in
hei
oos s
is
small
compa ed
wi h
ha
o ,
o
example,
pipis elles.
A
high
le el
o
dis u -
bance
by
humans
inside
oos s
p obably
causes
abandonmen ,
bu
in
gene al
P.
au i us
is
mo e
ole an
o
dis u bance
han
many
o he
species
which
li e
in
houses,
e.g.
Myo is
dauben onii
and
Myo is
na e e i
(Swi ,
unpublished).
A
some
oos s, long-ea ed
ba s
ha e
been
ound
o
e u n
ollowing
ex ensi e
eno a ions
o
he
house,
e lec ing
hei
loyal y
o
adi ional,
long-used
si es.
The
empe a u e
o
oo
spaces
was
ound
o
be
impo an
in
oos
selec ion
by
P.
au i-
us
(En wis le
e
al.,
1997).
Ba s
selec ed
wa me
buildings
-
he
mean
a e age
daily
em-
pe a u e
in
he
apex
o
oos s
was
17.9°C,
while
he
mean
a e age
daily
empe a u e
in
andomly
picked
houses
was
16.7°C.
The e
was
e idence
ha
ba s
we e
selec ing
no
only
wa me
buildings
bu
also
buildings
which
had
a
g ea e
capaci y
o
ap
hea .
The
p esence
o
sa king
may
be
connec ed
wi h
his,
since
i
imp o es
insula ion
and
so
educes
empe -
a u e
a ia ion.
How
ba s
seek
and
ind
sui able
oos s
is
no
clea .
Because
mos
a e
close
o
deciduous
woodland,
i
is
p obable
ha
buildings
a e
ound
and
in es iga ed
du ing
o aging
ligh s.
Smell
o
echoloca ion
may
be
used
(En wis le,
1994), o
i
is
possible
ha
ba s
may
be
a ac ed
by
he
p esence
o
conspeci ics.
Wha
is
clea ,
howe e ,
is
ha
long-ea ed
ba s
gain
a
numbe
o
ecological
ad an ages
by
selec ing
he
buildings
in
which
hey
oos .
Ba s
om
wa m
oos s
su ounded
by
la ge
a eas
o
deciduous
woodland
ha e
been
ound
o
g ow
as e ,
achie e
be e
body
condi ion
and
ha e
highe
su i al
a es
han
do
hose
om
less
a ou ed
oos s
(En wis le,
1994).
THERMOREGULATION
IN
ROOSTS
The
p e ious
chap e
has
shown
ha
he mo egula o y
beha iou
by
ep oducing
P.
au i-
us
is
complex.
Ba s
may
become
o pid
in
esponse
o
sho - e m,
occasional
ad e se
wea he
condi ions
and
hey
may
also
use
daily
o po
on
a
mo e
egula
basis
o
balance
high
ene gy
demand
a
ce ain
imes
du ing
summe ,
al hough
he e
is
e idence
ha
o po
is
incompa ible
wi h
ce ain
s ages
in
he
ep oduc i e
cycle,
e.g.
peak spe ma ogenesis
in
males
and
la e
p egnancy
in
emales.
By
selec ing
wa m
oo
spaces
as
oos s
and
adjus ing
94
Social
O ganiza ion
and
Beha iou
hei
beha iou
o
minimize
hea
loss
and
ene gy
expendi u e
du ing
imes
o
high
demand,
long-ea ed
ba s
a e
able
o
balance
hei
ene gy
budge
and
s ill
comple e
hei
ep oduc i e
cycle
wi hin
he
ime
a ailable.
Roos
Tempe a u e
The
di e ence
in
empe a u e
eco ded
be ween
P.
au i us
oos s
and
andomly
picked
houses
(En wis le
e al.,
1997)
was
no
due
o
hea ing
o
he
space
by
he
ba s
-
he
au ho s
epo ed
ha
empe a u e
p obes
placed
close
o
and
away
om
oos ing
ba s
showed
no
di e ences.
The
ba s
ac i ely
selec ed
wa me
houses
in
which
o
oos .
The
a e age
daily
empe a u e
o
nu se y
oos s
in
Sco land
anged
om
10.5°C
o
26.6°C,
while
he
mean
hou ly
empe a u e
anged
om
6.3°C
o
40.6°C
(En wis le,
1994).
Daily
a ia ion
in
empe a u e
was
simila
be ween
oos s
and
h oughou
summe .
En wis le
eco ded
ha
minimum
oos
empe a u e
occu ed
be ween
0500
and
0700
hou s,
and
oos s
hen
hea ed
apidly
o
a
maximum
empe a u e
in
mid
a e noon
(1200-1500
hou s)
be o e
declining
h ough
he
la e
a e noon
and
e ening.
Roos
empe a u e
was
in luenced
by
he
ai
empe a u e
ou side,
he
amoun
o
sola
adia ion
in
a
day
and
he
wind
speed.
Ba s
used
di e en
compa men s
wi hin
he
oos
space
o
selec
he
op imum
mic oclima e
a
any
ime
and
equen ly changed
compa men s
h ough
he
summe .
Coole
a eas
we e
gene ally
selec ed
in
June
and
Oc obe
(consis en
wi h
mo e
use
o
o po
a
hese
imes)
and
wa me
a eas
in
July
and
Augus ,
when
ep oduc i e
demands
on
ba s
esul ed
in
less
use
o
o po .
Indi iduals
we e
likely
o
eso
o
o po
o
conse e
ene gy
a
a
wide
ange
o
oos
empe a u es
below
abou
22°C
(En wis le,
1994).
Clus e ing
The
o ma ion
o
dense
g oups,
o
huddles,
by
indi iduals
may
be
used
ins ead
o
o po
o
conse e
ene gy,
o
bo h
may
be
used
oge he .
Clus e ing
is
pa icula ly
common
du ing
he
lac a ion
pe iod
in
many
species,
a
a
ime
when
ene gy
demand
is
high
and
ju eniles
ha e
o
main ain
a
ela i ely
high
body
empe a u e
in
o de
o
g ow.
T une
and
Slobodchiko
(1978)
ound
ha
pallid
ba s
(An ozouspallidas)
o med
clus e s
a
no mal
oos
empe a u es,
which
allowed
hem
o
educe
oxygen
consump ion
and
ene gy
expen-
di u e.
Ju eniles
we e
ound
a
he
cen e
( he
wa mes
pa )
o
he
clus e
mo e o en
han
would
ha e
been
expec ed
had
hei
posi ion
been
andom,
and
he
au ho s
p oposed
ha
he
adul s
we e
displaying
a
o m
o
ecip ocal
al uism
by
mu ually
allowing
each
o he 's
o sp ing
access
o
a
bene icial
posi ion.
C.
ownsendii
also
o med
densely
packed
clus e s
on
he
ceilings
o
he
ca es
whe e
hey
oos ed
du ing
he
lac a ion
pe iod
(Humph ey
and
Kunz,
1976).
Clus e ing
was
ound
by
En wis le
(1994)
o
be
e ec i e
in
educing
hea
loss
in
P.
au i-
us
-
labo a o y
expe imen s
showed
ha
a
clus e
o
ba s
in
a
small
space
aised
he ambi-
en
empe a u e
by
a ound
3°C.
In
oos s,
bo h
clus e ing
and
o po
we e
ound
o
be
used
du ing
he
s udy,
oge he
esul ing
in
signi ican
sa ings
in
ene gy
expendi u e
by
he
ba s.
The e
is
some
e idence
ha
P.
aus iacus
makes
less
use
o
clus e ing
beha iou
han
does
i s
congene ic.
S ebbings
(1970)
obse ed
ha
P.
aus iacus
did
no
o m
clus e s
du ing
summe
mon hs,
while
P.
au i us
in
he
same
a ic
did,
and Ho acek
(1975)
ound
ha
soli a y
indi iduals
in
a
oos
we e
much
mo e
likely
o
occu
among
P.
aus iacus.
This
Tempo a y
and
T ansi o y
Roos s
95
di e ence
may
be
connec ed
wi h
he
high
le el
o
in aspeci ic
agg ession
epo ed
in
g ey
long-ea ed
ba s
(S ebbings,
1970),
and
may
be
a
u he
eason
why
his
species seeks
wa me
a ics
han
does
P.
au i us
(Joo is,
1980)
and
why
i s
dis ibu ion
is
es ic ed
o
wa m,
lowland
egions.
Reac ion
o
Low
Tempe a u e
No mal
daily
luc ua ions
in
empe a u e
may
be
compensa ed
o
by
he
use
o
o po
and
clus e ing
wi hin
he
oos ,
bu
he e
a e
imes
du ing
sp ing
and
summe
when
se ious
ene gy
sho alls
a e
caused
by
ad e se
wea he
condi ions.
Ex ended
cold
spells
o
e y
we
wea he
cause
sho ages
in
insec
a ailabili y
and
may
mean
ha
o aging
is
unp oduc i e
o
e en
impossible.
A
such
imes,
P.
au i us
eac
by
empo a ily
abandoning
hei
nu se y
oos s
and
mo ing
o
al e na i e
oos s
in
he
a ea
(En wis le,
1994).
Such
oos s a e
no -
mally
in
ee
holes
o
a m
buildings
and
ha e
coole
mic oclima es
han
main
oos s.
A
adio
acking
s udy
(En wis le,
1994)
showed
ha
he e
was
a
s ong
co ela ion
be ween
he
occu ence
o
cold
nigh s
and
he
use
o
cool
al e na i e
oos s
by
P.
au i us.
Using
hese
oos s
allowed
ba s
o
d op
hei
body
empe a u e
u he
han
hey
could
in
he
wa me
main
oos s
and
so
conse e
mo e
ene gy
un il
condi ions
imp o ed.
Reac ion
o
High
Tempe a u e
S ebbings
(1966)
epo ed
ha
Pleco us
ba s
al e ed
hei
oos ing
beha iou when
he
ai
empe a u e
in
he
oos
exceeded
abou
40°C,
when
hey
hung
om
he
a e s
wi h
hei
wings
pa ly
sp ead.
Du ing
an
excep ionally
ho
spell
o
wea he
in
Sco land
du ing
Augus
1995,
I
isi ed
ou
P.
au i us
oos s
known
o
ha e
been
occupied
wi hin
he
p e-
ious
week.
Th ee
o
hem
we e
ound
o
be
emp y,
despi e
he
p esence
o
esh
d oppings.
In
he
ou h
I
ound
only
wo
ba s
(ou
o
a
colony
coun ed
he
p e ious
week
a
20)
and
bo h
o
hese
we e
hanging
om
a e s
wi h
hei
wings
sligh ly
sp ead,
in
he
posi ion
shown
in
Figu e
6.2.
The
empe a u e
a
he
oo
apex
was
39°C
a
1430
hou s
B i ish
Summe
Time
and
his
pa icula
oo
had
only
wo
compa men s,
bo h
o
which
we e
equally
ho .
I
he e o e
appea s
ha
oos s
can
become
oo
ho
o
long-ea ed
ba s.
In
an
a emp
o
lose
hea ,
hey
may
oos
away
om
he
apex
and
hang
in
open
ai
space
(Figu e
6.2),
bu
i
hey
s ill
o e hea
i
appea s
hey
ha e
o
abandon
he
a ic.
I
suspec
he
colonies
om
he
oos s
I
isi ed
had
empo a ily
mo ed
o
coole
al e na i e
oos s.
TEMPORARY
AND
TRANSITORY
ROOSTS
Tempo a y
Roos s
Some imes
e med
al e na i e
oos s,
hese
a e
used
by
colonies
o
sho
pe iods
when
con-
di ions
in
main
nu se y
oos s
a e
unsui able,
as
desc ibed
abo e.
En wis le
(1994)
ound
li le
e idence
ha
hey
a e
used
o
any
easons
o he
han
he mo egula o y
ones.
No
co -
ela ion
was
ound
be ween
hei
use
and
deg ee
o
dis u bance
a
he
main
oos ,
no
be ween
hei
occupa ion
and
changing
use
by
he
ba s
o
o aging
si es.
The e
was
also
no
la ge-scale
change
om
main
o
empo a y
oos s
such
as
migh
ha e
been
expec ed
i
he
%
J
s
*3
n
I
"5S
*3
c-
F
*3
;«-3s
£
ss
bO
^
.«
-S-.
s
5?
S
.^o
OS
-5!
1°
(N
-5
•5
o
S
u
~s
E
§
Nu se y
Colonies
97
pu pose
o
hei
use
was
o
p eda o
a oidance,
as
sugges ed
by
Taake
and
Hildenhagen
(1989).
Howe e ,
long-ea ed
ba s
may
also
swi ch
oos s
o
educe
le els
o
in es a ion
y
ec opa asi es
(see
Chap e
7)
and
his
was
no
in es iga ed
in
En wis le's
(1994)
s udy.
Baue o a
and
Zima
(1988)
ag eed
ha
al e na i e
oos s
we e
used
o
he mo egula o y
easons.
They
mis -ne ed
bo h
P.
au i us
and
P.
aus iacus
a
a
ca e
en ance
in
he
Czech
Republic
du ing
summe .
This
ca e
was
no
used
as
a
nu se y
oos
and
ew
long-ea ed
ba s
hibe na ed
he e.
The
highes
in ensi y
o
isi s
was
eco ded
in
la e
summe ,
and
he e
was
also
a
peak
o
ac i i y
a
he
ca e
in
Ap il-May.
These
a e
he
wo
imes
in
summe
when
o po
is
used
mos
by
he
ba s,
making
i
e y
likely
ha
he
cool
ca e
was
used
o
pe iods
o
o po
o
sa e
ene gy.
Gaisle
and
Hanak
(1969)
ound
ha
empo a y
oos s
we e
used
equally
by
bo h
P.
au i us
and
P.
aus iacus
in
summe ,
con i ming
ha ,
in
his
beha iou al
aspec ,
he e
is
no
di e ence
be ween
he
wo
species.
A ailable
e idence
seems
o
indica e
ha
al hough
sho - e m
use
is
made
o
cool
oos s
by
long-ea ed
ba s,
hey
a e
gene ally
ai h ul
o
one
main
nu se y
oos
all
summe .
In
his
espec
hey
di e
om
some
espe ilionid
specis
(e.g.
Nyc alus
noc ula),
which
change
oos s
equen ly
(Slui e
and
an
Hee d ,
1966).
T ansi o y
Roos s
These
a e oos s
used
du ing
mo emen s
be ween
hibe nacula
and
summe
oos s
and
back.
They
a e
occupied
spo adically
and
o
sho
pe iods, mainly
in
spnng
and
au umn,
al hough
males
may
also
use
hem
in
summe
(Ho acek,
1975).
Heise
and
Schmid
(1988)
main ained
ha
a e
he
b eak-up
o
nu se y
colonies
in
au umn
and
be ween
he
end
o
hibe na ion
and
he
ligh
o
nu se y
oos s
in
sp ing,
P.
au i us
li e
la gely
independen ly
o
one
ano he
and
seek
shel e
in
a
a ie y
o
oos s
including,
i
he
wea he
is
se e e,
cel-
la s,
ca es
and
mine
sha s
no mally
used
as
hibe nacula.
T ee
holes
and
ba
boxes
a e
also
used.
The e
seems
o
be
no
colony
cohesion
wi hin
hese
oos s.
As
well
as
shel e ing
ba s,
ansi o y
oos s a e
used
in
connec ion
wi h
ma ing
beha iou .
P.
au i us
ne ed
a
en ances
o
hibe nacula
du ing
la e
July
and
Augus
we e
all
ound
o
be
adul
o
imma-
u e
males
(Ho acek,
1975),
and
emales
had
all
depa ed
om
nu se y
oos s
by
la e
Augus
o
ea ly
Sep embe .
In
con as ,
P.
aus iacus
we e ne e
ne ed
a
hibe nacula
du -
ing
summe
and
ba s
we e
ound
in
nu se y
oos s
as
la e
as
Oc obe .
Ho acek
(1975)
sug-
ges ed
ha
male
P.
au i us
isi ed
he
ca es
as
po en ial
ma ing
si es
and
we e
hen
joined
he e
by
emales
o
ma ing in
au umn,
while
P.
aus iacus
ma ed
in
summe
oos s
and
so
did
no
isi
hibe nacula
a
his
ime.
Simila ly
in
sp ing,
P.
au i us
we e
ac i e
a ound
ca e
en ances
in
Ma ch
and
we e
p obably
using
he
en ance
egion
o
ma ing;
hey
did
no
mo e
in o
summe
oos s
un il
la e
Ap il.
P.
aus iacus,
which
a e
no
hough
o
ma e
in
sp ing,
did
no
display
hese
sp ing
ligh s
a ound
ca e
en ances.
NURSERY
COLONIES
Colony
Size
Long-ea ed
ba
colonies
a e
ypically
small
compa ed
wi h
hose
o
many
o he
species.
Those
o
P.
au i us
mos ly
numbe
om
10-20
adul s
(Speakman
e
al.,
1991a;
En wis le,
110
Popula ion
Biology
popula ion
size
o
P.
au i us
was
es ima ed
o
be
abou
370400
(Speakman,
1991a).
The
o al
B i ish
popula ion
o
P.
aus iacus
has
been
coun ed
a
abou
1300
(S ebbings
and
G i i hs,
1986)
and
is
con ined
o
an
a ea
o
a ound
1400
km2.
This
gi es
a
popula ion
densi y
o
he
species
o
abou
0.001
ba s
ha"1,
o
one
colony
o
20
ba s
in
e e y
15
10x10
km
squa es
-
a
densi y
o
abou
one- i een h
ha
o
P.
au i us
in
he
small
a ea
o
B i ain
in
which
P.
aus iacus
occu s.
SEX
RATIO
Benzal
(1991)
s udied
a
popula ion
o
P.
au i us
li ing
in
boxes
in
a
pine
plan a ion
in
cen al
Spain.
O e all,
he
numbe
o
males
in
he
boxes,
which
housed
mainly
nu se y
colonies,
was
high
and
he
sex
a io
did
no
di e
signi ican ly
om
1:1
(96
males:
101
emales).
Howe e ,
among
adul
ba s,
he
a io
was
biased
owa ds
emales
(49
adul
males:75
adul
emales).
Con e sely,
among
ju eniles,
emales
we e
always
less
nume -
ous
han
males
(47
ju enile
males:26
ju enile
emales).
In
his
popula ion,
he e o e,
i
appea ed
ha
he e
was
ei he
a
highe
p opo ion
o
males
a
bi h
o
a
highe
su i al
among
males
in
in ancy.
This
may
ha e
compensa ed
o
wha
appea ed
o
be
highe
mo ali y
in
males
han
in
emales
among
adul
ba s.
Benzal
(1991)
ound
ha
ju enile
males
we e
signi ican ly
ligh e
han
ju enile
emales
in
au umn
and
sugges ed
ha
his
implied
a
highe
mo ali y
among
males
du ing
hei
i s
win e ,
which
may
ha e
been
compensa ed
o
by
mo e
males
han
emales
being
bo n.
Boyd
and
S ebbings
(1989)
ound
ha ,
in
he
i s
5
yea s
o
hei
s udy,
he e
we e
signi ican ly
mo e
male
han
emale
ju eniles
in
he
popula ion
(59
males:
39
emales),
bu
ha
in
he
second
5
yea s
he
a io
did
no
di e
signi ican ly
om
1:1
(93
males:95
emales).
The
o e all
sex
a io
also
a ied
o e
he
cou se
o
he
s udy
—
he
p opo ion
o
males
inc eased
du ing
he
i s
5
yea s
and
hen
dec eased.
In
any
yea ,
a
g ea e
p opo ion
o
males
han
emales
we e
ju eniles,
bu
he
a io
o
ju enile
males
:
ju enile
emales
did
no
a y sig-
ni ican ly
om
1:
1
in
any
one
yea .
Thei
indings
hus
di e ed
om
hose
o
Benzal
(1991)
and
he
si ua ion
ega ding
he
sex
a io
a
bi h
is
no
clea .
The
ecen
inding
by
En wis le
e
al.
(in
p ess;
see
Chap e
5),
ha
he
no mally
accep ed
c i e ion
o
assessing
ma u i y
in
male
ba s
is
un eliable
o
P.
au i us,
may
explain
his
con usion.
Among
adul s
he e
does
appea
o
be
a
cons an
bias
owa ds
emales
(S ebbings,
1970;
Boyd
and
S ebbings,
1989;
Benzal,
1991),
sugges ing
ha
in e sexual
di e ences
in
he
ecology
and
beha iou
o
he
species
cause
a
highe
dea h
a e
among
males
han
emales.
FLIGHT
DISTANCES
Bo h
P.
au i us
and
P.
aus iacus
a e
conside ed
o
be
s a iona y
species
which
do
no
mo e
long
dis ances
be ween
summe
and
win e
oos s
(S elko ,
1969;
Gaisle
and
Hanak,
1969);
P.
au i us
is
known
o
occupy
he
same
oos s
all
yea
ound
in
some
a eas
(S elko ,
1969).
Reco ded
ligh s
o
ma ked
indi iduals
ha e
all
been
sho .
Ha ma a
(1987)
no ed
ha
bo h
species
mo ed
only
abou
500
m
be ween
oos s
in
woodland,
and
Pa k
e
al.
(in
p ess)
simila ly
eco ded
only
sho
mo emen s
(up
o
1.75
km)
by
P.
au i us
be ween
ba
Fligh
Dis ances
111
boxes
in
a
o es
a ea.
Gaisle
and
Hanak
(1969)
in es iga ed
ligh
dis ances
o
inged
indi-
iduals
o
bo h
long-ea ed
species
o e
20
yea s,
and
he
longes
ligh s
eco ded
we e
42
km
o
P.
au i us
and
62
km
o
P.
aus iacus-,
hese
we e
dis ances
be ween
summe
and
win e
oos s.
Long-ea ed
ba s
do,
howe e ,
occasionally
ly
u he
han
his.
A
g oup
o
P.
au i us,
appa en ly
lying
sou h-wes ,
landed
on
a
ship
70
km
om
he
Yo kshi e
coas
in
1948
and
ano he
was
ound
dead
on
a
ligh ship
50
km
eas
o
No olk
in
Oc obe
1968,
coinciden
wi h
an
in lux
o
bi ds
om
Scandina ia
(Co be ,
1970).
In
Sep embe
1995,
a
b own
long-ea ed
ba
was
ound
on
an
oil
pla o m
in
he
No h
Sea
150
miles
om
he
Sco ish
coas
(Hu son,
1996).
One
specimen
o
P.
aus iacus
was
ound
on
a
ligh ship
18
km
o
he
Sussex
coas
in
1969,
ollowed
by
one
ound
dead
on
he
coas
11
days
la e
(Co be ,
1971).
These
e en s
may
indica e
ha
bo h
species
unde ake
long-dis ance
ligh s
occasionally,
bu
hey
a e
mo e
likely
o
ha e
been
caused
by
ba s
being
acciden ally
blown
ou
o
sea
by
s ong
winds.
The e
is
no
e idence
ha
long-ea ed
ba s
o
ei he
species
make
o ganized,
egula
mig a o y
ligh s
o
any
g ea
dis ance,
unlike,
o
example,
N.
noc ula,
inged
indi iduals
o
which
ha e
been
eco ded
mig a ing
up
o
2400
km
in
Eu ope
(Hanak,
1966).
In
being
essen ially
s a iona y
species,
long-ea ed
ba s
esemble
o he s
in
he
pleco ine
g oup.
C.
ownsendii
is
also
a
ela i ely
seden a y
species
which
was
ound
o
ly
an
a e age
dis ance
o
only
11.6
km
om
nu se y
oos s
o
hibe nacula
(Humph ey
and
Kunz,
1976).
These
au ho s,
in
an
ex ensi e
s udy,
ound
ha
80%
o
inged
ba s
e u ned
o
he
same
hibe nacula
yea
a e
yea
and
ha
85.5%
o
inged
ba s
made
ligh s
o
less
han
1.6
km,
only
wo
ligh s
exceeded
8
km.
Homing
The
abili y
o
ba s
o
e u n
o
oos
si es
ollowing
ansloca ion
has
been
widely
epo ed,
al hough
how
hey
do
i
is
no
ully
unde s ood.
Vision
has
been
epo ed
o
be
an
impo -
an
ac o ,
since
blinded
Myo is
sodalis
showed
conside ably
less
homing
abili y
han
did
sigh ed
ones
(Ba bou
e
al.,
1966),
and
smell
also
has
an
e ec ,
bu
only
close
o
he
oos
(Da is,
1966).
Homing
abili y,
as
migh
be
expec ed,
has
been
shown
o
be
mos
de eloped
in
mig a o y
ba
species
(Wason,
1978)
and
he e o e
Pleco us
species
may
be
expec ed
o
be
ela i ely
poo
a
inding
hei
way
home.
In
ac ,
ew
in es iga ions
appea
o
ha e
been
ca ied
ou .
Rybe g
(1947)
showed
ha
P.
au i us
we e
able
o
ind
hei
way
back
o
oos s
om
abou
60
km
away
-
a
dis ance
in
he
same
ange
as
he
longes
na u al
ligh s
o
hese
ba s
(Gaisle
and
Hanak, 1969).
Also,
wo
cap i e
P.
au i us
which
escaped
in
Abe deen
we e
ound
o
ha e
e u ned
o
hei
oos
45
km
away
wi hin
10
days
(M.A.
Ande son,
unpublished).
Cock um
(1956)
ound
ha
only
ou
C.
ownsendii,
ou
o
a
o al
o
54
eleased,
had
e u ned
o
hei
oos
45
km
away
wi hin
2
days,
and
concluded
ha
hom-
ing
abili y
in
his
species
was
poo ly
de eloped.
This
appea s
o
con i m
ha
non-mig a o y
ba
species
a e
ela i ely
ine icien
a
homing.
Because
Pleco us
ba s
do
no
ly
long
dis-
ances
no mally,
hey
a e
likely
o
be
amilia
wi h
only
limi ed
geog aphic
a eas.
Thei
ligh
is
ene ge ically
expensi e
compa ed
wi h
ha
o
mos
o he
species
(Chap e
2)
and
hey
a oid
lying
ac oss
open
spaces.
They
hus
ha e
all
he
hallma ks
o
being
s a iona y
species
which
emain
in
a
limi ed
a ea
and
do
no
ha e
well-de eloped
homing
abili y,
al hough
hey
a e
able
o
ind
hei
way
back
o
oos s
o e
dis ances
o
up
o
60
km
wi h-
ou
appa en
di icul y.

112
Popula ion
Biology
PARASITES
Compa ed wi h
many
insec i o ous
ba s,
Pleco us
ha bou
ela i ely
ew
ec opa asi es
and
in es a ions on
indi iduals
a e
usually
ligh ,
pa icula ly
hose
on
adul s;
S ebbings
(1966)
epo ed
ha
ju eniles
equen ly
ha bou ed
highe
numbe s.
Low
in es a ions
o
ec opa -
asi es
a e
p obably
connec ed
wi h
small
colony
size
in
hese
ba s
—
pa asi es
such
as
mi es
and
leas
h i e
in
la ge
numbe s
on
hos s
which
li e
in
close
con ac
wi h
each
o he
and
be ween
which
ans e
is
easy.
Long-ea ed
ba s
li e
in
small
colonies
which
a e
o en
di ided
in o
se e al
g oups
wi hin
la ge
oo
spaces
and
so
he e
is
ela i ely
li le
con ac .
They
also
mo e
be ween
main
and
empo a y
oos s
(Chap e
6),
hus
u he
educing
he
oppo uni y
o
pa asi e
numbe s
o
inc ease.
Ec opa asi es
The
commones
o
hese
a e
leas
and
mi es.
Ve y
occasionally
a
nyc e ibiid
ly
is
epo ed
o
occu
on
long-ea ed
ba s,
al hough
nei he
Pleco us
species
is
conside ed
o
be
a
na u al
hos
o
hese
(Hu son,
1984).
K is o ik
(1982)
eco ded
one
specimen
o
Nyc e ibia
schmidlii
on
a
male
P.
au i us in
he
Slo ak
Republic;
he
conside ed
i s
p esence
o
be
casual
and
sugges ed
i
was
a
s ay
om
a
membe
o
ano he
ba
species
oos ing
in
he
icini y.
Nyc e ibiidae
(O de
Dip e a)
a e
minu e,
wingless
pa asi es
o
ba s;
hey
ha e
long
legs
and
ha e
become highly
specialized
on
hei
way
o
li e.
Fleas
Fleas
(O de
Siphonap e a)
a e
small,
wingless
insec s
which
a e
la e ally
la ened
in
shape
and
which
li e
pa asi ically
on
mammals
and
bi ds.
The
la ae
a e
non-pa asi ic.
Ischnopsyllidae
a e
a
amily
o
specialis
ba
leas
(Figu e
7.2),
and
se e al
species
o
hese
ha e
been
eco ded
on
long-ea ed
ba s.
Hai linge
and
Rup ech
(1992)
ound
Ischnopsyllus
hexa inus
on
P.
au i us
in
Poland
and
his
species
also
occu s
on
he
same
hos
in
B i ain
(S ebbings,
1977).
Nyc e idopsylla
longiceps
occu s
a ely
on
P.
au i us
in
win e
(Swi ,
1991).
Mi es
Mi es
(Class
A achnida;
O de
Aca i)
a e
e y
nume ous
and
widesp ead
and
many
a e
pa -
asi ic.
The
cephalo ho ax
and
abdomen
a e
used
so
ha
he
body
essen ially
consis s
o
one
piece
(Figu e
7.2)
and
mos
species
ha e
eigh
legs.
The
amily
mos
o en
ound
on
long-
ea ed
ba s
a e
Spinc u nidae,
and
he
species
Spinc u nixpleco ina
has
been
collec ed
om
P.
au i us
in
B i ain
(S ebbings,
1977),
Poland
(Hai linge
and
Rup ech ,
1992)
and
Ko ea
(Kim
and
Lee,
1990).
O he
species
o
mi e
eco ded
om
Pleco us
species
include
O ni honyssus
pipis elli,
Neomyobia
pleco ia
(Swi ,
1991)
and
Lep o ombidium
ussicum
(Hai linge
and
Rup ech ,
1992).
In e nal
Pa asi es
Ga dne
e
al.
(1987)
examined
blood
smea s
o
12
B i ish
ba
species,
including
P.
au i us
bu
no
P.
aus iacus,
o
pa asi es.
The
numbe
in
P.
au i us
was
low,
he
only
o ganism
ound
being
G ahamella
species.
This
is
an
in ae y h ocy ic
bac e ium
ound
in
a
wide
ange
o
mammals.
I
is
ansmi ed
by
leas
and
is
no
conside ed
o
be
ha m ul
o
he
hos .
P eda ion
113
FIG
7.2
A
lea
o
he
amily
Ischnopsyllidae
(a)
and
a
mi e
(b)
ypical
o
hose
ec opa asi ic
on
long-ea ed
ba s.
PREDATION
In
he
opics,
specialis
hawks
p ey
exclusi ely
on
ba s,
bu
in
Eu ope
mos
incidences
o
p eda ion
on
ba s
by
bi ds
appea
o
be
occasional
and
oppo unis ic.
Howe e ,
al hough
ba s a e
o
mino
impo ance
in
he
die
o
he
bi ds
which
ca ch
hem,
his
does
no
mean
ha
occasional
p eda ion
is
i ial
om
he
pe spec i e
o
he
ba ,
and
he
need
o
a oid
p eda ion
exe s
conside able
in luence
on
aspec s
o
hei
beha iou .
114
Popula ion
Biology
P eda ion
by
Bi ds
Co ids
ake
some
ba s
in
he
USA
and
A ica (Rose ea ,
1965)
and
gulls
ha e
been
obse ed
o
ca ch
one
in
B i ain
(Clee es,
1969),
bu
a
su ey
o
all
eco ds
showed
ha
p eda ion
on
ba s
by
hese
g oups
is
negligible
(Speakman,
1991a).
Hawks
and
alcons
ca ch
e y
ew,
e lec ing
he
diu nal
ac i i y
o
hese
bi ds.
The
ew
ba s
hey
do
ake
a e
likely
o
be
o
species
which
eme ge
ea ly
in
he
e ening,
be o e
wiligh
(e.g.
se o ines,
Ep esicus
se o inus).
Long-ea ed
ba s,
which
eme ge
much
la e ,
a e
e y
unlikely
o
encoun e
hem.
Howe e
long-ea ed
ba s
which
ly
in
dayligh
o
any
eason
(see
below)
un
a
much
highe
isk,
as
is
shown
in
a
epo
by
Simms
(1977).
A
male
and
a
emale
kes el,
Falco
innunculus,
we e
obse ed
pu suing
a
g oup
o
eigh
long-ea ed
ba s
du ing
an
ea ly
e ening
in
Augus
and
one
ba
was
caugh .
The e
is
e idence
om
ex ensi e
die a y
s udies
in
Eu ope
ha
ba s
egula ly
u n
up
in
he
die s
o
owls.
Tawny
(S ix
aluco)
and
ba n
(Ty o
alba)
owls
a e
he
mos
impo an
p eda o s,
and
long-ea ed
owls
(Asio
o us)
also
p ey
on
hem.
Ba n
owls
gene ally
o age
in
open
coun y
such
as
o e
ields,
bu
awny
owls
o age
p edominan ly
in
closed
habi a s,
mainly
woodland,
copses
and wooded
ga dens,
and
long-ea ed
owls
also
p e e
wooded
a eas,
pa icula ly
coni e ous
and
mixed
woodland
(Glue,
1970);
he
la e
wo
species
migh
he e o e
be
expec ed
o
p ey
o
a
g ea e
ex en
on
long-ea ed ba s,
which
also
p e-
e
such
habi a s.
Glue
(1970)
analysed
119
pelle
samples,
each
consis ing
o
be ween
22
and
52
indi idual
ba n
owl
pelle s.
Ou
o
a
o al
o
31
491
p ey
i ems
om
hese
pelle s,
he
ound
he
emains
o
11
ba
skele ons,
including
one
o
P.
au i us.
In
Poland,
Rup ech
(1979)
ound
ha
bo h
awny
and
ba n
owls
p eyed
on
ba s
and
ha
P.
au i us
was
one
o
he
commones
ba
species
in
he
pelle s
o
awny
owls.
The
awny
owl
seemed
o
be
a
mo e
specialis
p eda o
o
ba s
han
was
he
ba n
owl,
al hough
o
21
ba
species
which
occu
in
Poland,
emains
o
20
we e
ound
in
ba n
owl
pelle s;
hese
included
bo h
P.
au i us
and
P.
aus iacus.
In
a
la e
s udy
(Rup ech ,
1990),
his
au ho
es ima ed
ha
ba
emains
accoun ed
o
0.81%
o
he
con en s
o
awny
and
ba n
owl
pelle s
in
wes e n
Poland.
The
equency
o
di e en
ba
species
in
he
pelle s e lec ed
hei
ela i e
abundance
in
he
a ea,
indica ing
ha
owls
we e
no
being
selec i e.
This
was
con i med
in
ano he
Polish
s udy
(Lesinski,
1989),
which
ound
ha
P.
au i us
emains
occu ed
mo e
equen ly
han
did
hose
o
P.
aus iacus
in
ba n
owl
pelle s,
e lec ing
he
highe
local
abundance
o
P.
au i us.
Occasionally,
howe e ,
indi idual
owls
do
p ey
on
ba s
in
a
highly
oppo unis ic
way.
Jen zsch
(1992)
epo ed
ha
one
ba n
owl,
which
li ed
close
o
a
Myo is
myo is
oos
in
sou he n
Ge many,
appea ed
o
ha e
become
adep
a
ca ching
hese
ba s
—
9.1%
o
i s
die
consis ed
o
M.
myo is,
which
mus
ha e
had
a
conside able
ad e se
e ec
on
he
ba s'
pop-
ula ion
locally.
M.
myo is
is
he
species
mos
p eyed
upon
by
owls
in
Eu ope.
Pleco us
a e
gene ally
less
common
in
pelle s,
cons i u ing
only
0.61%
o
he
con en
o
pelle s
in
some
a eas
(Lipej
and
Gje kes,
1992).
Speakman
(1991a),
in
a
comp ehensi e
e iew
o
p eda-
ion
by
bi ds
on
ba s,
a emp ed
o
assess
he
impac
his
p eda ion
has
on
ba
popula ions
in
B i ain.
F om
published
eco ds,
he
es ima ed
he
o al
popula ion
o
bi ds
which
may
p ey
on
ba s,
he
o al
annual
ood
in ake
o
hese
bi ds
and
he
p opo ion
o
hei
die
which
was
likely
o
consis
o
ba s.
F om
hese
igu es
he
es ima ed
he
o al
numbe
o
ba s
ea en
in
a
yea .
This
included
8783
ba s
ea en
by
ba n
owls,
168
850
by
awny
owls
and
10
205
by
long-ea ed
owls.
Nex ,
Speakman
es ima ed
he
o al
popula ion
o
ba s
in
B i ain
and
hei
o e all
mo ali y
a es
om
published
epo s.
F om
hese
he
calcula ed
P eda ion
115
he
o al
numbe
o
ba s
which
die
in
he
coun y
in
a
yea .
This
igu e
was
abou
1.82
mil-
lion,
o
which
201
415
we e
calcula ed
o
be
killed
by
bi ds,
indica ing
ha
a ian
p eda o s
accoun ed
o
a ound
11.1%
o
he
o al
annual
mo ali y
o
all
ba s
in
B i ain.
The
UK
popula ion
o
P.
aus iacus,
whose
oos s a e
ew
and
whose
ange
is
e y
lim-
i ed
in
B i ain,
has
been
es ima ed
a
a ound
1500
indi iduals
(S ebbings
and
G i i hs,
1986)
and
i s
annual
mo ali y
a e
is
0.33
(S ebbings,
1970).
The
o al
popula ion
o
P.
au i us
is
no
known,
bu
i s
popula ion
densi y
(Boyd
and
S ebbings,
1989;
Speakman
e
al.,
1991a)
was
calcula ed
by
Speakman
(1991a)
o
be
abou
one- en h
ha
o
P.
pipis el-
lus
and
i s
ange
in
B i ain
o
be
abou
80%
o
he
a ea
inhabi ed
by
pipis elles.
He
he e-
o e
es ima ed
he
popula ion
o
be
abou
8%
o
ha
o
pipis elles,
which
is
a ound
4.631
million
(Walsh
e
al.,
1987),
o
abou
370
000.
The
annual
mo ali y
a e
o
P.
au i us
is
0.242
(S ebbings,
1977).
F om
hese
igu es,
i
i
is
assumed
ha
bi ds
p ey
equally
on
all
ba
species,
a ian
p eda ion
accoun s
o
abou
9950
b own
and
55
g ey
long-ea ed
ba s
e e y
yea .
Speakman's
(1991a)
igu e
o
11.1%
o
he
impac
o
p eda ion
by
bi ds
on
ba
mo al-
i y
is
su p isingly
high,
gi en
he
low
incidence
o
ba s
in
he
die s
o
hese
bi ds.
I
is
also
s ange
ha
p eda ion
on
ba s
is
no
epo ed
mo e
o en,
pa icula ly
ha
by
diu nal
bi ds
such
as
kes els.
Speakman
(1991a)
es ima ed
ha
8406
ba s
a e
killed
by
kes els
e e y
yea ,
which
implies
ha
23
a e
a acked
e e y
day
by
hese
bi ds,
bu
kes els
chasing
ba s
is
s ill
a
a ely
eco ded
e en .
The
eason
o his
is
he
la ge
popula ion
o
kes els
in
he
UK.
In
o de
o
23
ba s
o
be
killed
e e y
day,
an
indi idual
kes el
would
only
need
o
ca ch
one
once
in
20
yea s.
High
owl
popula ions
in
he
UK
p obably
also
accoun
o
he
ela i ely
low
p opo ion
o
ba s
in
hei
die s.
Because
owl
popula ions
a e
na ionally
high,
hei
a e
o
p eda ion
on
ba s
need
only
be
ela i ely
low
o
p oduce
a
no iceable
e ec .
I
canno
he e o e
be
a gued
ha
p eda ion
is
oo
low
in
his
coun y
o
p oduce
an
e ec
on
he
beha iou
o
ba s
such
as
Pleco us
species,
and
i
is
highly
p obable
ha
beha iou
such
as
la e
eme gence,
selec ion
o
oos s
close
o
woodland
and
he
use
o
lyways
a e
connec ed
wi h p eda o
a oidance.
P eda ion
by
Mammals
The
mos
signi ican
mammalian
p eda o
o
long-ea ed
ba s
in
B i ain
and
Eu ope
is
almos
ce ainly
he
domes ic
ca ,
Felis
ca us.
In
he
Ne he lands,
eco ds
o
p eda ion
on
ba s
by
ca s
a e
almos
ou
imes
mo e
equen
han
hose
o
p eda ion
on
hem
by
owls
(Bekke
and
Mos e ,
1990),
and
ba s
which
oos
in
houses
a e
especially
ulne able
o
ca
p eda ion.
Unlike
bi ds
o
p ey,
domes ic
ca s
li e
in
close
p oximi y
o
humans
and
ha e
no
ea
o
app oaching
human
dwellings
-
hey
a e
hus
a
mo e
likely
han
owls
o
a ack
ba s
in
he
immedia e
icini y
o
he
oos .
They
a e
agile
and
adep
a
climbing
and
can
he e o e
ge
in o
a
posi ion
o
a ack
ba s
as
hey
eme ge.
They
may
e en
be
able
o
ge
in o
some
a ics
whe e
long-ea ed
ba s
oos
and
a ack
hem
he e.
B uijn
(1990)
pho og aphed
he
ac i i ies
o
a
ca
which
was
suspec ed
o
killing
a
numbe
o
ba s
(mainly
P.
pipis ellus
and
P.
au i us)
oos ing
in
a
owe .
The
ca
was
ilmed
en e ing
he
owe
h ough
a
en i-
la ion
opening,
climbing
80
s eps
o
he
lo
and
hen
a acking
he
oos ing
ba s.
Be o e
i s
ac i i ies
we e
cu ailed
by
ixing
a
g ill
o
he
en ila ion
opening,
i
had
killed
a
leas
50
ba s.
Many
had
been
egu gi a ed
o
ejec ed
unea en,
a
ea u e
common
in
ca
p eda ion.
Like
many
insec i o es
such
as
sh ews
and
moles, ba s a e
appa en ly
unpala able
o
ca s,
116
Popula ion
Biology
bu
his
does
no
de e
hem
om
killing
la ge
numbe s,
pe haps
in
he
hope
o
inding
one
which
as es
be e !
Anecdo al
e idence
sugges s
ha
mos
long-ea ed
ba
p eda ion
by
ca s
occu s
as
he
ba s
eme ge
in
he
e ening.
Ca s
climb
on
o
window
sills
o
oo s
and
cap u e
he
ba s
as
hey
d op
h ough
he
exi
hole
om
he
oos .
Many
ca s
become
e y
skilled
a
his
ac i i y
and
can
ha e
a
se ious
e ec
on
a
colony.
Many
o
he
long-ea ed
ba s
b ough
o
me
as
casual-
ies
ha e
been
escued
om
ca s,
and
hose
ound
a e
likely
o
o m
only
a
small
p opo ion
o
hose
caugh .
Slow- lying
ba s
such
as
Pleco us
species
a e
pa icula ly
ulne able.
Bagley
and
Jacobs
(1985)
eco ded
high
mo ali y
in
a
colony
o
he
ela ed
species
C.
ownsendii
i ginianus
due
o
a
domes ic
ca
ca ching
hem
on
eme gence.
The
colony's
ma e ni y
oos
was
in
a
ca e
whose
en ance
was
g illed,
and
he
ca
had
become
adep
a
ca ching
ba s
as
hey
lew
h ough
he
g ill.
DAYLIGHT
FLYING
Ba s
a e
almos
exclusi ely
noc u nal,
bu
obse a ions
and
epo s
indica e
ha
hey
do
occasionally
ly
du ing
dayligh .
Because
dayligh
lying
is
so
unusual,
na u alis s
end
o
no ice
and
eco d
ins ances
o
i ,
and
Speakman
(1990)
conduc ed
a
su ey
in
which
he
sough
as
many
obse a ions
as
possible
om
all
o e
B i ain
in
an
a emp
o
disco e
how
p e alen
dayligh
lying
was
and
wha
he
easons
we e
o
i s
occu ence.
A
o al
o
420
eco ds
we e
ecei ed,
abou
one- hi d
o
which
we e
o
ligh s
du ing
win e
(Oc obe -Ma ch).
In
he
emaining
wo- hi ds,
a
peak
o
ac i i y
occu ed
du ing
Ap il;
ac i i y
in
bo h
summe
and
win e
was
g ea es
in
he
middle
o
he
day,
be ween
1200
and
1600
hou s.
The
numbe
o
ba s
in ol ed
in
each
sigh ing
a ied
be ween
1
and
200,
bu
a ound
90%
we e
o
single
indi iduals.
Speakman
concluded
om
da a
in
he
su ey
ha
he
main
pu pose
o
dayligh
ligh s
in
summe
was
o
compensa e
o
ene gy
de ici s
caused
by
inadequa e
ood
in ake
du ing
noc u nal
o aging
—
i
poo
wea he
educed
ba s'
abili y
o
eed,
possibly
o
se e al
nigh s,
hey
eme ged
du ing
dayligh
as
an
eme -
gency
measu e.
The
p e alence
o
ligh s
in
Ap il
e lec ed
he
lowe
insec
popula ion
a
ha
ime
and
he
ba s'
subsequen
need
o
supplemen
nigh
eeding.
This
was
suppo ed
by
he
inding
ha
ba s
a
high
la i ude
we e
mo e
likely
o
ly
by
day
han
we e
hose
u -
he
sou h,
since
sho
nigh s
u he
added
o
hei
inabili y
o
mee
ene gy
demand
in
cold
o
we
condi ions.
In
win e ,
dayligh
lying
occu ed
because
pe iods
o
a ousal
om
hibe na ion
(Chap e
8)
did
no
always
coincide
wi h
pe iods
o
da kness.
I
ba s
a oused
du ing
dayligh
and
condi ions
we e
sui able
o
o aging,
hey
eme ged
bo h
o
eed
and
o
d ink.
O
he
420
eco ds o
ba s
lying
in
dayligh ,
16
we e
o
P.
au i us
and
one
o
P.
aus ia-
cus,
al hough
some
o
he
153
uniden i ied
ba s
may
also
ha e
been
Pleco us
species.
Speakman
(1990)
obse ed
ha
P.
au i us
hus
lew
in
dayligh
e y
in equen ly,
al hough
he
numbe
o
ligh s
made
by
his
species
(16)
and
he
numbe
made
by
P.
pipis ellus
(154)
appea
o
be
in
app oxima ely
he
same
p opo ion
as
hei
popula ions
in
B i ain,
since
he
P.
au i us
popula ion
was
es ima ed
by
Speakman
(1991a)
o
be
8%
o
ha
o
P.
pipis ellus.
The e
was
he e o e
no
e idence
ha
long-ea ed
ba s
we e
less
inclined
o
ly
by
day
han
any
o he
species,
despi e
hei
no mal
habi
o
o aging
la e
a
nigh
a he
han
in
wiligh .
O e all,
howe e ,
he
su ey
showed
ha
dayligh
ligh s
we e
a e,
and
Dayligh
Flying
117
Speakman
(1990)
calcula ed
ha
ba s
o
all
species
we e
abou
100
imes
less
likely
o
ly
by
day
han
by
nigh .
Since
peak
insec
a ailabili y
occu s
in
mid
o
la e
a e noon
in
empe a e
egions
(Rydell
and
Speakman,
1995)
and
since
ba s
equen ly
ha e
o
eso
o
ene gy
sa ing
beha iou
such
as
o po
and
clus e ing
du ing
ep oduc ion
when
demand
is
high,
Speakman
(1991b)
nex
sough
easons
o
explain
why
dayligh
lying
does
no
occu
mo e
equen ly.
Fou
al e na i e
hypo heses
we e
examined:
•
Di ec
compe i ion
wi h
insec i o ous
bi ds
(e.g.
house
ma ins)
may
educe
he
amoun
o
ood
ba s
could
acqui e
by
day.
•
Mobbing
by
bi ds
(e.g.
c ows)
which
a e
nei he
p eda o s
no
compe i o s
may
educe
eeding
e iciency.
•
O e hea ing
caused
by
he
incidence
o
sunligh
on
ba s'
da k,
uninsula ed
wings
may
mean
hey
a e
unable
o
dissipa e
hea
gene a ed
in
ligh
(Speakman
and
Hays,
1992).
•
Dayligh
lying
may
expose
ba s
o
an
unaccep ably
high
isk
o
a ack
by
diu nal
a ian
p eda o s.
Speakman's
(1990)
su ey
showed
ha
encoun e s
be ween
dayligh
lying
ba s
and
mobbing
bi ds
we e
a e
and
i ial
and
could
no
explain
he
in equency
o
dayligh
ly-
ing.
In e ac ions
wi h
a ian ae ial
insec i o es
we e
also
in equen
( hey
occu ed
in
only
1.4%
o
eco ds),
and
a
sepa a e
s udy
(Ve nie ,
1990)
ound
no
e idence
o
compe i ion
wi h
hi undine
bi ds.
Fossil
e idence
also
makes
he
compe i ion
hypo hesis
unlikely
-
by
he
ea ly
o
mid
Eocene,
mic ochi op e an
ba s
we e
al eady
specialized
o
echoloca ion
bu
passe ine
bi ds,
including
Hi udinidae,
did
no
appea
be o e
he
Oligocene
(Rydell
and
Speakman,
1995);
ba s
he e o e
had
no
po en ial
compe i o s
du ing
he
Eocene.
Howe e ,
he
ossil
his o y
o
owls
da es
back
o
he
Palaeocene
and
ha
o
o he
p eda o y
bi ds
such
as
hawks
o
he
ea ly
Eocene
(Rydell
and
Speakman,
1995)
and
hus
he e
we e
po en ial
p eda o s
o
ba s
a
he
ime
hei
noc u nal
habi s
we e
e ol ing.
In
o de
o
in es iga e
he
p eda ion
and
o e hea ing
hypo heses,
Speakman
(1991b)
conduc ed
a
u he
analysis
on
da a
om
his
dayligh
lying
su ey
and
also
in es iga ed
lying
beha iou
in
cap i e
P.
pipis ellus
and
P.
au i us.
Expe imen s
we e
pe o med
o
es ablish
he
e ec
o
ligh
on
he
lying
beha iou
o
he
ba s
and
whe he
he e
was
any
e ec
o
adian
hea
on
body
empe a u e.
Indi idual
ba s
o
bo h
species
lew
o
a
signi -
ican ly
g ea e
pe cen age
o
ime
in
he
da k
han
in
he
ligh ;
he e
was
no
di e ence
be ween
he
species
and
bo h
a oided
lying
in
he
ligh .
The e
was
also
no
signi ican
di -
e ence
in
ei he
species
be ween
he
body
empe a u es
o
ba s
measu ed
a
he
end
o
ligh s
in
he
da k
and
in
he
ligh ,
indica ing
ha
a oidance
o
ligh
was
no
media ed
by
o e hea ing.
O e hea ing
was
hus
no
he
p oximal
cue
which
caused
he
ba s'
a oidance
o
lying
in
he
ligh ,
al hough
i
could
no
be
uled
ou
as
an
ul ima e
ac o
in luencing
he
eason
o
he
a oidance
o
dayligh
lying.
The
su ey
(Speakman,
1990)
also
showed
ha
ba s
did
no
pa icula ly
a oid
lying
on
ho ,
sunny
days,
sugges ing
ha ,
a
leas
in
he
UK,
o e hea ing
does
no
ha e
a
signi ican
e ec .
The
mos
likely
ac o
which
causes
ba s
o
be
noc u nal
hus
appea s
o
be
p eda ion,
and
he
esul s
o
he
su ey
(Speakman,
1991b)
suppo
his.
O
he
420
epo s
o
day-
ligh
ligh s,
3.1%
included
obse a ions
o
in e ac ions
wi h
ca s
o
wi h
a ian
p eda o s
including
kes els,
me lins
(Falco
columba ius),
spa owhawks
(Accipi e
nisus)
and
118
Popula ion
Biology
Time
spen
lying
in
dayligh
(h)
FIG
7.3
P obabili y
o
su i ing
p eda ion
a emp s
agains
ime
spen
lying
in
day-
ligh
o
ba s
in
he
UK
( ep oduced
wi h
pe mission
om
Speakman,
1991b).
black-headed
gulls
(La us
idibundus).
In
a
leas
1.2%
o
he
420
epo s,
he
ba
was
killed.
Speakman
es ima ed,
om
da a
in
he
su ey,
ha
on
a e age
a
ba
lying
in
dayligh
would
be
a ally
a acked
abou
once
in
e e y
14.3
h
o
ligh .
A
ba
lying
in
dayligh
o
64
h
would
ha e
a
99%
chance
o
being
killed
(Figu e
7.3).
This
is
abou
100
imes
he
es i-
ma ed
isk
o
dea h
by
noc u nal
p eda ion,
and
s ongly
suppo s
he
hypo hesis
ha
a oidance
o
diu nal
p eda o s
is
he
eason
why
ba s
a e
e y
la gely
noc u nal.
CHAPTER
8
Hibe na ion
B
ATS
li ing
in
empe a e
egions
ace
an
annual
p oblem
o
ood
sho age,
since
ew
a h opods
a e
a ailable
du ing
win e .
All
animals
which
depend
on
seasonal
ood
sou ces
ha e
he
same
p oblem
and
se e al
solu ions
ha e
e ol ed.
Animals
may
swi ch
ood
sou ces
empo a ily,
e.g.
he
bank
ole,
Cle h ionomys
gla eolus,
may
ea
dead
lea es
in
mid
win e
when
i s
no mal
die
o
seeds,
ui s
and
lea es
o
woody
plan s
is
no
a ailable
(Wa s,
1968).
Howe e ,
his
op ion
is
no
open
o
ba s
because
no
app op ia e
al e na i e
oods
a e
a ailable
o
hem.
Mig a ion
o
lowe
la i udes
is
common
among
bi d
species
and
has
also
e ol ed
among
some
ba s.
In
Eu ope,
species
such
as
P.
pipis ellus
and
TV.
noc ula
mig a e
o e
la ge
dis ances,
pa icula ly
in
no he n
pa s
o
hei
ange
(S elko ,
1969).
The
hi d
s a egy
is
hibe na ion,
an
op ion
open
only
o
animals,
such
as
empe a e
zone
ba s,
which
can
become
o pid.
Ba s
some imes
use
bo h
s a egies
—
pipis elles
and
noc-
ules,
when
hey
mig a e,
do
so
om
summe
o
win e
oos s,
which
a e
usually
u he
sou h,
and
he e
hey
hibe na e.
S a iona y
species
such
as
long-ea ed
ba s,
whose
summe
and
win e
oos s a e
close
o
each
o he ,
ely
en i ely
on
hibe na ion
as
a
s a egy
o
su i e
cold
win e s
and
se e ely
cu ailed
ood
supplies.
HETEROTHERMY,
TORPOR
AND
HIBERNATION
He e o he my
is
a
specialized
o m
o
endo he my
ound
mainly
among
small,
empe a e
zone
mammals.
A
he e o he mic
animal
is
able
o
a y
i s
body
empe a u e
bu ,
unlike
ec o he mic
animals
such
as
ep iles,
he e o he ms
a e
able
o
con ol
he
all
in
body
em-
pe a u e
and
also
o
es o e
high
body
empe a u e
independen ly
o
ambien
empe a u e.
They
do
his
using
hea
gene a ed
by
mobiliza ion
o
a
s o ed
in
he
body.
To po
is
de ined
as
a
s a e
in
which
an
animal
allows
i s
body
empe a u e
o
all
below
i s
ac i e
endo he mic
le el.
The
d op
in
empe a u e
is
con olled
and,
a
low
ambien
empe a u e,
can
be
main ained
wi hin
na ow
limi s
by
me abolic
p ocesses.
Hibe na ion
is
an
ex ended
o m
o
o po
which
may
las
o
days
o
weeks
and
which
occu s
in
esponse
o
a
p olonged
all
in
ambien
empe a u e
and
a
educ ion
in
ood
supply.
I
is
no ,
howe e ,
con inuous
o po
which
las s
all
win e .
Ransome
(1971)
p oposed
ha
i
di e s
om
sho - e m
o po
used
by
ba s
du ing
summe
only
in
how
o en
a ousal
occu s.
In
summe ,
a
ba
a ouses
equen ly
(hence
he
e m
'Tagesschla le ha gie'
(day-
sleep
le ha gy)
used
by
Eisen au ,
1937),
while
in
win e
a ousals
may
occu
days
o
weeks
apa
(B ack
and
Twen e,
1983;
Twen e
e
al.,
1983;
Thomas,
1995a).
119
120
Hibe na ion
Al ingham
(1996)
lis ed
ou
ea u es
o
o po
in
an
a emp
o
de ine
i
in
physiologi-
cal
e ms:
•
I
in ol es
a
con olled
educ ion
o
body
empe a u e,
ypically
o
wi hin
1—2°C
o
ambien
empe a u e.
•
Hea
a e,
oxygen
consump ion,
b ea hing
a e
and
me abolic
a e
all
all
as
he
body
empe a u e
d ops.
•
Pe iphe al
asocons ic ion
occu s
and,
a
ex emes
o
empe a u e,
blood
low
is
es ic ed
o
a
ew
i al
o gans
o
much
o
he
ime.
•
The
o pid
animal
is
able
o
a ouse
spon aneously,
independen ly
o
ambien
empe -
a u e.
Timing
o
Hibe na ion
S ebbings
(1970)
eco ded
ha
P.
aus iacus
in
sou he n
England
began
o
hibe na e
ea lie
han
P.
au i us;
whe eas
mos
P.
aus iacus
om
a
colony
we e
hibe na ing
by
la e
Oc obe
o
ea ly
No embe ,
P.
au i us
om
he
same
oos
we e
s ill
ac i e
in
ea ly
No embe
and
he
whole
colony
was
no
in
hibe na ion
be o e
he
end
o
No embe .
In
cen al
Eu ope,
Hu ka
(1971)
simila ly
ound
ha
P.
aus iacus
began
o
hibe na e
in
Oc obe .
In
Poland,
Ha ma a
(1973),
who
did
no
dis inguish
be ween
he
wo
species,
ound
ha
Pleco us
ba s
i s
appea ed
in
hibe nacula
a
he
beginning
o
No embe
and
s ayed
he e
un il
mid
Ma ch.
S ebbings
(1970)
ound
ha
emale
P.
aus iacus
began
o
eme ge
om
hibe na ion
in
la e
Ma ch
and
by
ea ly
Ap il,
all
ba s
o
his
species
we e
ac i e.
P.
au i us
we e
all
ac i e
by
mid
Ap il.
In
Eu ope,
ba s
o
bo h
species
ended
hibe na ion
in
la e
Ma ch
(Ha ma a,
1973),
al hough
Ho acek
(1975)
ound
ha
a
ew
P.
au i us
emained
in
hibe nacula
un il
ea ly
Ap il.
Fa
Accumula ion
P io
o
Hibe na ion
All
hibe na ing
ba s
accumula e
a
du ing
la e
summe
and
au umn
and
his
is
used
as
an
ene gy
s o e
o
win e .
The
weigh
gain
can
be
conside able
-
Ransome
(1990)
eco ded
ha
he
a es
indi iduals
o
Rhinolophus
e umequinum
in
his
s udies
gained
75-80%
o
hei
lean
weigh
du ing
Oc obe ,
al hough
weigh
gain,
e en
among
ba s
o
he
same
species,
was
e y
a iable.
Hale
(1980)
caugh
a
o al
o
12
P.
au i us
in
mis
ne s
du ing
a
s udy
o
bi d
mig a ion
in
England
and
eco ded
he
weigh s
o
hese
be o e
eleasing
hem.
On
26
Augus ,
he
a e age
weigh
o
males
among
ne ed
indi iduals
was
6.50
g
(«
=
3)
and
ha
o
emales
7.65
g
(«
=
2).
On
13
Oc obe ,
he
a e age
weigh s
o
ba s
was
9.25
g
(n
=
2)
o
males
and
10.82
g
(n=
5)
o
emales,
which
indica ed
ha
he
a e age
weigh
o
ba s
sampled
had
inc eased
by
41%
o e
a
pe iod
o
7
weeks.
This
gain
is
high
compa ed
wi h
mos
ba
species,
in
which
p e-hibe nal
weigh
gain
is
a ound
25-30%
(Humph ey
and
Kunz,
1976).
In
Spain,
weigh
gain
in
P.
au i us
a
he
same
ime
o
yea
was
ound
o
be
less;
indi idual
adul
emales
sampled
weighed
an
a e age
o
9.0
g
in
July
and
11.2
g
in
Oc obe ,
ep esen ing
a
gain
o
25%
(Benzal,
1991).
The
di e ence
be ween
he
ba s
in
he
wo
s udies
was
ha
hose
in
Spain
had
a
highe
minimum
weigh
han
hose
in
England.
Ei he
he
ue
minimum
weigh
in
he
Spanish
ba s
was
missed
o
else
long-ea ed
ba s
a
his
lowe
la i ude
ne e
lose
as
much
a
and
so
hei
weigh
ne e
alls
below
abou
9
g.
Si e
Selec ion
121
High
p e-hibe nal
weigh
gain
was
also
ound
in
he
pleco ine
species
C.
ownsendii,
in
which
a
accumula ion
accoun ed
o
50%
o
he
weigh
o
males
and
57%
o
ha
o
emales
immedia ely
be o e
hibe na ion
(Humph ey
and
Kunz,
1976).
These
au ho s
sug-
ges ed
ha
la ge
weigh
gain
may
ha e
compensa ed
o
he
high
equency
wi h
which
he
species
mo ed
hibe na ion
si es
du ing
win e
and
he
consequen
high
a e
a
which
hey
used
up
ene gy.
Funakoshi
and
Uchida
(1982)
sugges ed
ha
p e-hibe nal
weigh
inc ease
in
ba s
depends
on
an
endogenous
ci cannual
hy hm
which
is
independen
o
he
iming
o
he
ep oduc i e
cycle,
since
la e
bi hs
du ing
a
summe
had
no
e ec
on
p e-hibe nal
weigh
gain
among
espe ilionid
ba s
in
hei
s udy.
They
also
sugges ed
ha
he
mos
likely
Zei gebe ,
o
exogenous
iming
ac o ,
was
ambien
empe a u e.
Weigh
Loss
Du ing
Hibe na ion
S ebbings
(1970)
es ima ed
weigh
loss
du ing
hibe na ion
in
sou he n
England
o
be
22%
in
P.
au i us
and
29%
in
P.
aus iacus.
G ey
long-ea ed
ba s
a oused
mo e
o en
du ing
win-
e
and
changed
hibe na ion
si es
mo e
equen ly,
which
accoun ed
o
hei highe
weigh
loss.
S ebbings
a ibu ed
his
o
unsui able
hibe na ing
condi ions
o
his
species
in
B i ain,
which
o ced
ba s
o
a ouse
o en
o
eed.
He
sugges ed
ha
P.
aus iacus
a e
con-
ined
o
he
ex eme
sou h
o
he
coun y
because
his
is
he
only
pa
whe e
he
numbe
o
insec s
lying
in
win e
is
high
enough
o
allow
hem
o
do
so.
SITE
SELECTION
Hibe na ion
Si es
Pleco us
species
hibe na e
in
ca es,
bo h
na u al
ones
and
man-made
s uc u es
such
as
mines,
qua ies
and
unde g ound
o i ica ions.
They
a e
also
among
he
Eu opean
ba s
mos
likely
o
be
ound
in
buildings,
mainly
cella s,
and
hey
may
spend
a
leas
pa
o
he
win e
in
ee
holes.
In
gene al,
hey
choose
ela i ely
cool
si es
and
mo e
be ween
hibe -
nacula
mo e
equen ly
han
do
mos
Eu opean
ba s.
Daan
and
Wiche s
(1968)
ound
ha
Pleco us
species
spen
less
ime
in
win e
in
a
sys-
em
o
a i icial
limes one
ca es
han
did
any
o
he
o he
eigh
species
which
used
he
ca es.
Thei
s ay
was
con ined
o
a
pe iod
in
he
middle
o
win e ,
and
i
was
p esumed
ha
hey
spen
he
es
o
he ime
in
coole
si es
such
as
ees.
Howe e ,
in
Poland
long-ea ed
ba s
we e
ound
o
spend
mo e
ime
in
unde g ound
si es,
pa icula ly
in
a
sys em
o
o i ica-
ions
(Be na d
e
al.,
1991).
In
Sweden,
whe e
he e
a e
ew
na u al
ca es,
P.
au i us
ha e
been
eco ded
hibe na ing
in
mines
and
limes one
qua ies,
and
34%
o
he
ba s
in
one
mine
in
he
sou h
o
he
coun y
we e
o
his
species
(Ge ell,
1980-81).
Whe e
he
wo
Pleco us
species
hibe na e
in
he
same
si es
in
cen al
Eu ope,
P.
aus iacus
has
been
ound
o
selec
sligh ly
wa me
mic oclima es
(Hu ka,
1971;
Ho acek,
1975).
When
long-ea ed
ba s
hibe na e
in
buildings,
cella s
a e
he
commones
si es
o
hem,
al hough
one
specimen
o
P.
aus iacus
was
ound
in
Decembe
in
a
disused
b ick
kiln
in
he
Ne he lands
(Glas,
1982).
In
Ge many,
a
soli a y
P.
aus iacus
hibe na ed
in
a
cas le
cel-
la
nea
Be lin
(Haensel
and
Na e,
1993),
whe e
Haensel
(1994)
epo ed
ha
P.
au i us

122
Hibe na ion
was
he
mos
nume ous
ba
species
hibe na ing
in
unde g ound
cella s
o
ba ns
and
houses.
Cella s
a e
also
common
si es
o
ind
P.
aus iacus
in
he
Czech
Republic
(Ho acek,
1975)
and
in
Ge many
(Haensel,
1994).
Bo h
Ge ell
(1980—81)
and
Rydell
(1989c)
epo ed
ex ensi e
use
o
s one-buil
cella s
by
P.
au i us
in
Sweden.
These
cella s,
which
a e
e y
common
in
he
sou h
o
he
coun y,
a e
de ached
om
houses
and
ha e
a
aul ed
oo .
They
we e
adi ionally
used
o
s o ing
po a oes,
bu
nowadays
a e
equen ly
emp y.
Ge ell
(1980-81)
ound
ha
P.
au i us
was
he
mos
nume ous
ba
species
in
hem
in
win-
e .
Rydell
(1989c)
sea ched
s one
cella s
in
many
u al
a eas
in
Sweden
o
signs
o
ba s
and
ound
ha
63%
o
in ac
cella s,
bu
only
11%
o
de elic
ones,
con ained
such
signs.
Rydell
concluded
ha
hese
si es
we e
impo an
hibe nacula
o
P.
au i us.
The
condi ion
o
he
cella
was
ob iously
impo an ,
and
hus
he e
was
a
need
o
p ese e
hem
in
good
epai ,
al hough
ba s
we e
no
de e ed
by
he
cella s
ul illing
hei
no mal
unc ion
o
s o -
ing
po a oes.
Occupa ion
was
ound
o
be
highes
in
hose
con aining
many
c e ices,
which
we e
used
as
oos
si es.
Changing
Si es
by
Ba s
Long-ea ed
ba s
change
hei
hibe na ion
si es
equen ly
du ing
win e .
Daan
(1970)
used
au oma ic
lash
pho og aphy
in
a
ca e
sys em
in
he
Ne he lands
o
moni o
si e-changing
ac i i y
by
ba s.
He
ound
ha
Pleco us
species
made
mo e
ligh s
inside
he
ca es
and
changed
hibe na ion
si es
mo e
o en
han
did
indi iduals
o
se e al
Myo is
species
which
also
hibe na ed he e.
Bezem
e
al.
(1964)
s udied
mo emen s
o
P.
au i us
in
a
complex
sys em
o
a i icial
limes one
ca es
and
ound
ha ,
s a is ically,
his
species
showed
a
p e -
e ence
o
oos ing
in
he
cool
ou e
egion
o
he
ca es,
up
o
50
m
om
he
en ance.
Howe e ,
in
se e ely
os y
wea he ,
hey
mig a ed
o
he
wa me
inne
egion,
mo e
han
50
m
om
he
ou side.
This
indica ed
ha
long-ea ed
ba s
changed
si es
wi hin
hibe nac-
ula
in
esponse
o
changes
in
ambien
empe a u e.
Kuipe s
and
Daan
(1970)
also
s udied
in e nal
mig a ion
in
he
same
ca e
sys em.
Thei
s udy
in ol ed
a
numbe
o
species
including
g ey
and
b own
long-ea ed
ba s.
Pa s
o
he
ca es
whe e
ela i ely
wa m
ai
could
low
in o
he
en ance
a ea
om
ou side
we e
a oided
by
long-ea ed
ba s
ea ly
in
he
win e ;
hey
hibe na ed
in
he
deepe
a eas,
which
we e
coole .
As
he
en ance
a ea
cooled
down
wi h
cold
ai
lowing
om
ou side,
hey
mo ed
in o
i
om
he
inne
a eas.
Howe e ,
in
one
pa
o
he
sys em,
di e en
ai
low
mean
ha
he
en ance
a ea
was
he
cooles
pa
all
win e ;
in
his
sec ion,
long-ea ed
ba s
spen
all
hei
hibe na ion
pe iod
in
he
en ance
a ea.
Tempe a u e
All
eco ds
a ailable
show
ha
long-ea ed
ba s
p e e
lowe
ambien
empe a u es
a
which
o
hibe na e
han
do
mos
o he
Eu opean
species.
Daan
and
Wiche s
(1968)
epo ed
ha
he
a e age
empe a u e
in
a
limes one
ca e
close
o
hibe na ing
Pleco us
species
anged
om
0-5.3°C,
which
was
lowe
han
ha
close
o
any
o
he
o he
eigh
species
in
he
same
ca e.
Ha ma a
(1969,
1973)
measu ed
he
p e e ed
hibe na ing
empe a u es
o
se e al
Eu opean
species,
bo h
unde
na u al
condi ions
and
in
he
labo a o y.
In
cap i i y,
P.
au i us
chose
o
hibe na e
a
empe a u es
o
1-8°C,
wi h
mos
indi iduals
selec ing
6°C
(Ha ma a,
1969).
Tempe a u es
selec ed
in
he
labo a o y
we e
gene ally
sligh ly
highe
Si e
Selec ion
123
han
hose
selec ed
in
he
wild.
In
ano he
s udy
unde
na u al
condi ions,
long-ea ed
ba s
(no
dis inc ion
made
be ween
species)
hibe na ed
a
empe a u es
o
be ween
-3
and
11
°C,
wi h
mos
indi iduals
choosing
a eas
a
a ound
7°C
(Ha ma a,
1973).
Gaisle
(1970)
used
a
na u al
si ua ion
o
show
ha
Pleco us
p e e
o
hibe na e
a
low
empe a u es.
A
ca e
in
he
Czech
Republic
was
di ided
in o
wo
pa s
by
a
wooden
wall
and
one
sec ion
was
con-
side ably
wa me
han
he
o he ;
a
numbe
o
long-ea ed
ba s
egula ly
hibe na ed
in
he
coole
sec ion.
One
au umn,
he
wall
was
b oken
down,
allowing
wa m
ai
in o
he
coole
pa
o
he
ca e,
and
he
ollowing
win e
no
Pleco us
ba s
hibe na ed
he e.
They
e u ned
g adually
a e
he
wall
was
es o ed.
Nagel
and
Nagel
(1991)
epo ed
ha
P.
au i us
in
Ge many
hibe na ed
a
a
mean
ambien
empe a u e
o
4°C.
Ha ma a
(1969)
ound
ha
P.
au i us
emained
ac i e
a
empe a u es
as
low
as
1°C,
which may
explain
why
hey
equi e
low
empe a u es
o
hibe na ion
and
why
hey
may
be
ound
in
summe
oos s
well
in o
au umn
(Chap e
6).
Ha ma a
(1969)
epo ed
ha
he
only
o he
Eu opean
species
which
showed
a
simila
p e e ence
o
low
hibe na ion
empe a u es
was
Ba bas ella
ba -
bas ellus,
ano he
membe
o
he
pleco ine
g oup
(see
Chap e
1).
O he
pleco ine
ba s
(e.g.
C.
ownsendii;
Humph ey
and
Kunz,
1976)
ha e
also
been
ound
o
p e e
empe a u es
in
he
same ange.
Bo h
long-ea ed
ba s
and
ba bas elles
a e
also
able
o
ole a e
lowe
ex emes
o
empe a u e
han
a e
o he
species
in
Eu ope
-
Abelence
e
al.
(1956)
epo ed
ha
P.
au i us
could
su i e
a
empe a u es
as
low
as
-7.5°C
and
B.
ba bas ellus
se e al
deg ees
below
his.
Whe e
da a
a e
a ailable
o
b own
and
g ey
long-ea ed
ba s
sepa a ely,
hey
indica e
ha
he
anges
o
empe a u es
a
which
hey
hibe na e
a e
simila
—
Ho acek
(1975)
ound
P.
aus iacus
hibe na ing
a
2-9°C
-
bu
ha
P.
aus iacus
in
gene al
p e e
sligh ly
wa me
hibe nacula.
Humidi y
Rela i e
humidi y
in
ba
hibe nacula
is
gene ally
high
(Al ingham, 1996)
and
is
likely
o
be
highe
in
ca es
han
in
buildings.
Speakman
and
Racey
(1989)
p oposed
ha
hibe na -
ing
ba s a e
mo e
a
isk
om
dehyd a ion
han
om
s a a ion
and
ha
he
main
pu pose
o
mos
win e
ligh s
is
o
d ink
a he
han
o
eed.
By
selec ing
humid
hibe nacula,
ba s
a e
able
o
educe
wa e
loss
while
hey
a e
o pid
and
so
ha e
o
a ouse
less
o en.
Daan
and
Wiche s
(1968)
ound
he
ela i e
humidi y
o
limes one
ca es
a ound
hibe na ing
Pleco us
o
be
95-100%.
Simila
high
humidi y
was
ound
in
ca es
con aining
hibe na ing
long-
ea ed
ba s
in
Russia
(75-100%;
Lesinski,
1986)
and
Poland
( ange
55-100%,
a e age
84%;
Bogdanowicz
and
U banczyk,
1983).
Al i ude
The e
a e
a
numbe
o
epo s om
Eu ope
ha
long-ea ed
ba s
use
hibe nacula
a
high
al i ude.
In
sou he n
Ge many,
Nagel
e
al.
(1983—4)
ound
P.
au i us
and
a
ew
P.
aus i-
acus
in
ca es
a
700-800
m
abo e
sea
le el,
while
e y
ew
we e
ound
below
his
al i ude
and
negligible
numbe s
hibe na ed
below
500
m.
In
summe ,
howe e ,
almos
all
ba s
in
he
a ea
we e
ound
a
lowe
al i udes.
I
was
he e o e
appa en
ha
long-ea ed
ba s
mo ed
in o
he
moun ains
in
win e ,
p obably
because
empe a u es
in
ca es
he e
we e
lowe .
A
ca e
a
800
m
abo e
sea
le el
in
he
a ea
was
ound
o
be
2—3°C
coole
han
one
a
300
m
124
Hibe na ion
abo e
sea
le el.
Nagel
and
Nagel
(1991)
simila ly
ound
ha
P.
au i us
in
Ge many
p e-
e ed
coole
si es
a
high
al i ude
in
which
o
hibe na e.
Si es
Chosen
wi hin
Hibe nacula
Bezem
e
al.
(1964)
desc ibed
h ee
possible
posi ions
o
ba s
wi hin
hibe nacula:
•
hey
may
hang
eely
om
a
ho izon al
su ace
such
as
a
ceiling
•
hey
may
hang
in
con ac
wi h
a
wall
•
hey
may
be
in
c e ices,
ei he
deep
ones
such
as
bo e
holes
in
qua ies
o
shallow
ones
such
as
inden a ions
in
ock.
The
nine
species
hibe na ing
in
a
la ge
sys em
o
a i icial
ca es
in
he
Ne he lands
we e
anked
by
hese
au ho s
in
o de
o
hei
inclina ion
o
hibe na e
in
c e ices,
om
Rhinolophus
hipposide os,
which
was
always
ound
hanging
eely
om
he
oo ,
o
P.
pip-
is ellus,
which
in a iably
hibe na ed
deep
in
c e ices:
R.
hipposide os,
Myo is
ema gina us,
M.
myo is,
M.
dasycneme,
M.
na e e i,
M.
mys acinus
and
P.
au i us,
M.
dauben onii,
P.
pip-
is ellus.
P.
au i us
was
hus
in
he
middle
o
he
ange
and
was
usually
ound
ei he
in
con-
ac
wi h
a
wall
o
in
c e ices.
They
used
bo h
shallow
and
deepe
issu es.
In
he
same
ca es,
howe e ,
Daan
and
Wiche s
(1968)
always
ound
long-ea ed
ba s
(no
dis inc ion
was
made
be ween
he
wo
species)
in
c e ices
-
hey
we e
ne e
on
walls.
Ho acek
(1975)
ound
di e ences
in
he
hibe na ion
si es
used
by
P.
au i us
and
P.
aus-
iacus
in
he
Czech
Republic.
Wi hin
a
limes one
qua y,
P.
au i us
we e
usually
ound
in
la ge
galle ies,
and
indi iduals
hibe na ed
in
c e ices
o
bo e
holes,
mos ly
a
dep h
(Figu e
8.1).
They
appea ed
o
p e e
si es
whe e
ambien
empe a u e
luc ua ed
li le
(hence
hei
p e e ence
o
deepe
c e ices)
and
whe e
he
ange
was
2-7°C.
P.
aus iacus
hibe na ed
in
wa me
pa s
o
he
qua y
(2-9°C),
o en
in
smalle
galle ies
and
in
si es
subjec ed
o
con-
side able
empe a u e
changes.
They
we e
ne e
ound
deep
in
c e ices,
bu
p e e ed
o
be
on
walls
o
a
he
en ances
o
c e ices
(Figu e
8.2).
Ho acek
(1975)
also
equen ly
ound
P.
aus iacus
in
cella s
du ing
win e ,
and
concluded
ha
hey
we e
mo e
likely
han
P.
au i-
us
o
hibe na e
in
synan h opic
si es.
In
gene al,
bo h
species
o
long-ea ed
ba s
hibe na e
singly
o
in
e y
small
g oups
o
wo
o
h ee
indi iduals
(Ha ma a,
1973).
When
hey
a e
in
such
small
g oups,
indi idual
ba s
do
no
o m
di ec
body
con ac
wi h
each
o he ,
and
huddles
o
clus e s
ha e
no
been
epo ed.
Pleco us
ba s
a e,
occasionally,
ound
in
la ge
g oups
wi h
o he
species
which
ha e
simila
he mal
p e e ences
(Bogdanowicz,
1983),
bu
hey
do
no
o m
la ge
clus e s.
AROUSAL
A ousal
is
he
p ocess
by
which
ba s
ac i ely
eme ge
om
a
s a e
o
o po .
They
may
become
ac i e
in
esponse
o
dis u bance
in
he
hibe naculum,
bu
a ousal
also
occu s
ou-
inely
unde
na u al
condi ions.
The
p ocess
in ol es
he
use
o
b own
adipose
issue,
which
is
ound
in
he
neck
and
scapula
egion
in
empe a u e
zone
ba s.
Obse a ions
o
long-ea ed
ba s
ac i e
in
win e
unde
na u al
condi ions
(e.g.
Daan,
1970)
sugges
hey
FIG
8.1
Hibe na ion
si es
commonly
occupied
by
P.
au i us.
Win e
Ac i i y
127
a ouse
ela i ely
equen ly
compa ed
wi h
mos
o he
species.
Daan
(1973)
ound
ha
among
ba s
o
h ee
species
o
Myo is,
a ousal
occu ed
on
a e age
e e y
20
days
du ing
mid
win e .
In
Oc obe
and
in
ea ly
sp ing
ac i i y
was
clea ly
noc u nal
bu
du ing
he
middle
o
win e
ba s
a oused
a
any
ime
o
day,
and
dayligh
ligh s
occu ed.
This
pa e n
was
consis en
wi h
a ousal
being
con olled
by
a
ee- unning
endogenous
hy hm.
In
au umn
and
sp ing,
wi h
sho e
pe iods
o
o po ,
he
hy hm
was
en ained
o
a ousals
a
dusk.
A
simila
si ua ion
was
ound
o
exis
in
he
Ame ican
species
Myo is
luci ugus
by
Thomas
(1995a),
using
adio
eleme y
in
o de
o
a oid
dis u bing
ba s
and
so
causing
non-na u al
a ousals.
Ba s
a oused
e e y
15-20
days
a
andom
imes
-
he e
was
no
endency
o
become
ac i e
a
dusk.
Thomas
conside ed
ha
he
equency
o
a ousal
was
mos
p obably
go e ned
by
wa e
loss
in
ba s
and
hei
consequen
need
o
become
ac i e
in o de
o
d ink.
F equency
o
a ousal
in
P.
au i us,
and
he
ac o s
go e ning
i ,
ha e
been
s udied
unde
semi-na u al
condi ions
(Hays
e
al.,
1992)
and
a e
discussed
below.
WINTER
ACTIVITY
Ba s
which
a ouse om
o po
may
hen
lea e
he
hibe naculum
and
ly
ac i ely.
S ebbings
(1966,
1970)
epo ed
in e mi en
ac i i y
among
bo h
P.
au i us
and
P.
aus iacus
h oughou
win e ,
including
one
ligh
a
midday
on
a
Janua y
day
when
he
ai
empe a-
u e
was
below
0°C
and
snow
was
lying.
A e y
(1985)
in es iga ed
win e
ac i i y
in
P.
pip-
is ellus
in
sou he n
England
and
ound
ha
ba s
eme ged
du ing
all
win e
mon hs,
ha
ac i i y
le els
we e
highe
on
wa m,
calm
nigh s
and
ha
le el
o
ac i i y
was
co ela ed
wi h
day ime
empe a u e
and
no
wi h
he
empe a u e
he
nigh
be o e.
The
p ima y
unc ion
o
win e
ligh s
is
no
clea .
Feeding
may
ake
place
du ing
hem
(Ransome,
1968;
A e y,
1985)
and
bo h
A e y
(1985)
and
B igham
(1987)
sugges ed
ha
he
need
o
eed
was
he
mos
impo an
s imulus.
Howe e ,
Speakman
and
Racey
(1989)
concluded,
om
a
s udy
on
cap i e
P.
pipis ellus,
ha
hibe na ing
ba s
would
die
o
dehy-
d a ion
be o e
hey
s a ed
and
ha
win e
ligh s
we e
ini ia ed
by
he
need
o
d ink.
They
sugges ed
ha
ba s
which
ed
du ing
win e
ligh s
did
so
in o de
o
co e
he
ene ge ic
cos
o
he
ligh ,
bu
ha
his
was
no
hei
main
pu pose.
Because
in
a
na u al
si ua ion
ba s
almos
always
ea
and
d ink
du ing
ligh s,
i
is
di icul
o
sepa a e
he
wo
ac i i ies.
In
an
a emp
o
cla i y
he
si ua ion
in
P.
au i us,
Hays
e
al.
(1992)
in es iga ed
he
daily
ood
and
wa e
consump ion
and
he
indi idual
p obabili y
o
eme gence
in
a
cap i e
colony.
The
ba s
we e
kep
in
a
ee- ligh
enclosu e
in
which
hey
we e
exposed
o
na u al
pho ope iod
and
en i onmen al
empe a u e
and
whe e
hey
had
unlimi ed
access
o
d ink-
ing
wa e
bu
only
pe iodic
access
o
ood.
They
hibe na ed
in
a
small
wooden
box
inside
he
enclosu e.
Ac i i y
was
moni o ed
by wo
Dopple
ada
uni s
in e aced
o
a
mic o-
compu e .
Du ing
he
expe imen ,
which
ook
place
be ween
Janua y
and
Ma ch,
ba s
we e
subjec ed
o
11
days
wi h
access
o
wa e
only,
hen
14
days
wi h
access
o
ood
and
wa e ,
11
days
wi h
wa e
only
and
inally
15
days
wi h
ood
and
wa e .
The
ba s
we e
weighed
a
he
beginning
and
end
o
each
pe iod,
bu
o he wise
we e
comple ely
undis u bed.
Physical
condi ions
in
he
oos
box
we e
wi hin
he
ange
ound
in
na u al
hibe nacula
-
he
mean
FIG
8.2
Hibe na ion
si es
commonly
occupied
by
P.
aus iacus.
128
Hibe na ion
ela i e
humidi y
was
82%
and
he
empe a u e
was
3.4-12.4°C. The
esul s
o
he
s udy
showed
ha
he
p obabili y
ha
ba s
would
eme ge
inc eased
when
he e
was
ood
a ailable
and
when
i
was
wa me .
Food
consump ion
was
signi ican ly
and
posi i ely
co ela ed
wi h
he
mean
empe a u e
a
nigh
in
he
ligh
enclosu e;
ba s
we e
mo e
ac i e
and
a e
and
d ank
mo e
on
wa me
nigh s.
When
ood
was
no
a ailable,
ba s
eme ged
on
14
ou
o
22
nigh s
and
when
ood
was
a ailable,
hey
eme ged
on
28
ou
o
29
nigh s.
Eme gence
hus
occu ed
equen ly
e en
when
ood
was
no
a ailable,
sugges ing
ha
in
his
species
win e
ligh s
may
be
almos
a
daily
occu ence
excep
in
e y
cold
wea he .
The
au ho s
p oposed
ha ,
in
P.
au i us,
win e
ligh s
may
no
be
ini ia ed
by
he
onse
o
ei he
dehy-
d a ion
o
s a a ion.
Ra he ,
a
ypical
win e
empe a u es,
ba s
may
ly
whene e
poss-
ible,
almos
daily,
in o de
o
ensu e
ha
nei he
wa e
no
ood
ese es
app oach
c i ical
le els.
P olonged
bou s
o
o po
(mo e
han
a
ew
days)
only
occu
i
ambien
empe a u e
d ops
o
less
han
abou
4°C.
Hays
e
al.'s
(1992)
s udy
also
showed
ha
win e
ac i i y
in
P.
au i us
was
s ic ly
noc-
u nal.
The
ime
o
ini ial
ac i i y
was
s ongly
co ela ed
wi h
he
ime
o
sunse
(Figu e
8.3),
wi h
i s
eme gence
occu ing
a
mean
in e al
o
64.4
min
a e
sunse .
I
he e o e
appea s
ha ,
in
his
species,
equen
and
egula
ac i i y
in
win e
p e en s
he
endogenous
hy hm
con olling
a ousal
om
becoming
ee- unning
and
main ains
en ainmen
o
he
ligh -da k
cycle.
Dayligh
lying
does
occasionally
occu
(Speakman,
1990),
bu
p obably
only
in
esponse
o
ex ended
cold
spells
and
subsequen
long
pe iods
o
o po .
The
eason
why
P.
au i us
lies
mo e
equen ly
in
win e
han
many
o he
species
may
be
connec ed
wi h
i s
habi
o
gleaning
much
o
i s
ood
(Chap e
3).
While
mos
ba
species
will
only
be
able
o
eed
on
nigh s
when
he
ai
empe a u e
is
abo e
he
h eshold
o
insec
ligh
(which
a ies
o
di e en
species
bu
is
in
he
ange
o
5—
10°C;
Taylo ,
1963;
Rydell,
1989a),
long-ea ed
ba s a e
able
o
glean
insec s
c awling
on
ege a ion
a
lowe
empe a u es.
Thei
o aging
me hod
also
allows
hem
o
eed
on
diapausing
insec s
Da e
FIG
8.3
The
ime
o
i s
eme gence
o
P.
au i us
hibe na ing
in
semi-na u al
con-
di ions,
plo ed
agains
da e
du ing
Janua y
and
Feb ua y.
The
line
ep esen s
he
ime
o sunse
a
57
°N
( ep oduced
wi h
pe mission
om
Hays
e
al.,
1992).
E ec s
o
Dis u bance
on
Hibe na ing
Ba s
129
(Roe ,
1969).
The e
is
hus
mo e
accessible
ood
a ailable
o
long-ea ed
ba s
han
o
non-
gleaning
species,
e en
on
colde
nigh s.
Howe e ,
hey
a e
mo e
ac i e
on
wa me -nigh s
and
du ing
mild
spells
in
win e
hey
spend
less
ime
in
o po
and
eed
o
longe
(
al.,
1992).
They
also
d ink
mo e
on
nigh s
when
ood
is
a ailable,
sugges ing
a
mos
he
wa e
d unk
is
o
balance
he
wa e
loss
incu ed
as
a
esul
o
s aying
ac i e
and
eeding.
EFFECTS
OF
DISTURBANCE
ON
HIBERNATING
BATS
Dis u bance
inside
hibe nacula
has
long
been
conside ed
a
cause
o
he
declineo
ba
popula ions.
Besides
delibe a e
des uc ion
o
colonies,
inad e en
dis u bance
by
ou is
and
speleologis s
and
by
conse a ionis s
and
biologis s
moni o ing
popula ion
is
houg
o
ha e
had
a
de imen al
e ec .
Dis u bance
causes
unna u al
a ousal
in
hibe na ing
ba s,
which
hen
use
up
ene gy
esou ces
and
may
in
consequence
un
ou
o
ene gy
and
die
be o e
he
end
o
win e .
La ge-scale
handling
and
inging
o
hibe na ing
ba s
was
den
-
ied
as
ha ing
a
majo
ad e se
e ec
on
ba
numbe s
(Pun ,
1970),
and
Gaisle
e
al.
(1981)
demons a ed
ha
popula ion
educ ions
in
some
ca e-dwelling
species
we e
a
di ec
co
-
sequence
o
hei
own
esea ch
ac i i ies.
S ebbings
(1966,
1969)
no ed
ha
handling
and
weighing
a
long-ea ed
ba
caused
i
o
lose
0.18
g
pe
day,
which
he
es ima ed
o
be
3-4
imes
he
no mal
daily
weigh
loss
o
a
o pid
ba .
The e o e a
ba handled
six
imes
du ing
a
win e
could
lose
3-4
weeks'
wo h
o
hibe na ion
ime
due
o
loss
o
a mid
his
cou
make
he
di e ence
be ween
su i ing
and
no
doing
so.
K zanowski
(1961)
epo
simila
weigh
losses
due
o
handling
in
ca e-dwelling
ba s
in
Poland.
As
a
esul
o
indings
such
as
hese,
guidelines
we e
d awn
up
ecommending
ha
isi
s
o
hibe nacula
should
be
es ic ed
o
2-3
pe
win e
and
ba s
should
no
be
handled,
p
o-
og aphed
using
lash
o
exposed
o
b igh
o chligh ,
excessi e
noise
o
smoking
(Hu son
and
Micklebu gh,
1988).
Howe e ,
he
ela i e
e ec s
o
di e en
snmuh
we e
unknown,
so
in
a
se ies
o
labo a o y
expe imen s
on
cap i e
ba s,
Speakman
e
al
(1991b)
assessed
he
equency
and
magni ude
o
a ousals
p o oked
by
a ious
ac ile
and
non- ac ile
s im-
uli.
Ba s
o
six
species
including
P.
au i us
we e
kep
in
a
cool
ligh
oom
whe e
ood
and
wa e
we e
cons an ly
a ailable.
Indi iduals
we e
weighed
and
hen
placed
in
a
espi a o y
chambe
a
a
empe a u e
o
1-5°C
h ough
which
cool
ai
lowed;
hey
we e
hen
le
undis u bed
un il
hey
had
become
o pid.
A e
his,
ene gy
expendi u e
was
calcula ed
om
oxygen
consump ion.
Inside
he
chambe ,
ba s
we e
subjec ed
o
non- ac ile
s imuli
which
included
o chligh ,
pho og aphic
lash,
sound,
human
speech
and
bu s s
o
wa m
ai
o
7-10
min.
Tac ile
s imulus
was
p o ided
by
handling
he
ba
in
he
chambe
h ough
a
ubbe
glo e
i ed
o
he
oo .
A
posi i e
esponse
was
de ined
as
a
signi ican
inc ease
in
ene gy
expendi u e
abo e
he
p e-s imula ion
o pid
a e,
and
esponse
was
moni o ed
un il
he
ba
e u ned
o
o po .
The
suscep ibili y
o
ba s
o
all
i e
non- ac i
e
s imuli
was
low.
P.
au i us
esponded
o
only
wo
o
hem
-
one
indi idual
o
pho og aphic
lash
and
ano he
o
wa m
ai .
In
bo h
cases,
he
deg ee
o
dis u bance
was
low
and
he
ime
be o e
he
ba
again
became
o pid
was
sho .
Howe e ,
ac ile
s imulus
elici ed
a
esponse
in
e e y
case
and
he
equency
o
esponse
was
signi ican ly
highe
han
ha
o
non- ac-
ile
s imuli.
The
ene gy
expendi u e
pe
esponse
was
also
signi ican ly
highe
(Figu e
8.4);
he
au ho s
calcula ed
ha
he
a e age
ex a
ene gy
expended
by
b own
long-ea ed
ba s
du ing
a
esponse
o
a
non- ac ile
s imulus
was
8
J,
bu
ha
expended
in
esponse
o
a
142
Long-Ea ed
Ba s
and
Humans
eeding
include
Rhinolophus
species
(Jones
e
al.,
1995).
These
ba s
all
ly
ela i ely slowly
and
manoeu ably
and
use
ei he
e y
long,
high- equency,
na ow-band
calls
o
b oad-
band
echoloca ion
pulses
wi h
as
epe i ion
a e;
bo h
ypes
o
call
a e
sui able
o
o ag-
ing
in
clu e ed
si ua ions
(Baagoe,
1987).
I
is
hus
possible
ha
P.
au i us
does
no
o age
ound
s ee
lamps
ei he
because
i s
me hod
o
o aging
is
unsui able o
ca ching
insec s
lying
as
in
open
si ua ions
o
because
i
a oids
he
ligh
in
o de
o
educe
expo-
su e
o
p eda o s.
Baue o a
(1982)
obse ed
P.
aus iacus
o aging
ound
s ee
lamps
in
he
Czech
Republic.
Al hough
la ge
numbe s
o
small
mo hs
swa med
ound
a
ligh ,
he
ba
igno ed
hem
and,
on
eigh
occasions,
pu sued
and
caugh
la ge
mo hs
in
as
ligh .
G ey
long-
ea ed
ba s
hus
appea
o
di e
om
b own
ones
in
his
aspec
o
beha iou ;
hey
do
exploi
s ee lamps
and
use
he
oppo uni y
p o ided
o
pu sue
la ge
mo hs
in
ee
ligh .
A oiding
lying
in
he
ligh
does
no
seem
o
be
impo an
o
his
species.
Ano he
obse a ion
om
Eu ope
con uses
he
issue
u he ;
Ba a aud
(1990)
s udied
a
mixed
colony
o
P.
au i us
and
P.
aus iacus
and
du ing
his
s udy
he
analysed
pe ch
emains
om
unde
a
willow
ee
si -
ua ed
di ec ly
benea h
a
s ee
lamp.
O e
60%
o
he
emains
we e
om
la ae
and
adul s
o
mo h
species
whose
la ae
eed
on
willow
lea es,
and
i
hus
appea ed
ha
a
leas
one
Pleco us
ba
exploi ed
insec s
in
his
ee.
The
au ho
did
no
s a e
which
species
used
he
pe ch
—
i
i
was
P.
au i us,
hen
he
p esence
o
he
s ee ligh
does
no
seem
o
ha e
de e ed
i
om
eeding
he e,
mainly
by
gleaning.
I
he
ba
was
an
indi idual
o
P.
aus i-
acus,
hen
i
mus
ha e
gleaned
a
mo e
han
his
species
is
epo ed
o
do,
in
o de
o
ha e
caugh
he
la ae.
I
is
possible
ha
P.
au i us
a e
no
de e ed
by
ligh
and
ha
hey
do
glean
a ound
s ee
lamps
on
occasions
when
he
oppo uni y
a ises.
I
his
is
he
case,
hen
hei
no mal
a oidance
o
s ee
lamps
is
due
o
hei
inabili y
o
exploi
a
si ua ion
in
which
mo hs
a e
lying
as
and
in
he
open,
a he
han
o
a
need
o
a oid
p eda ion.
Fu he
esea ch
is
needed.
Implica ions
o
Long-ea ed
Ba s
The
limi ed
e idence
so
a
a ailable
indica es
ha
P.
aus iacus
is
able
o
exploi
s ee
lamps
o
o aging
bu
ha
P.
au i us,
in
gene al,
is
no .
Illumina ed
oads
will
p obably
con inue
o
be
a ailable
in
he
u u e,
unlike
many
o he
ba
habi a s
which
a e
becom-
ing
ewe ,
and
his
will
bene i
P.
aus iacus.
The e
has,
howe e ,
been
a
endency
in
ecen
yea s
in
Eu ope
o
eplace
me cu y
apou
ligh s
wi h
sodium
ones,
which
use
less
ene gy
and
which
do
no
equi e
he
p ocessing
o
handling
o
poisonous
me cu y
(Rydell
and
Racey,
1995),
and
sodium
lamps
ha e
been
shown
o
a ac
a
ewe
insec s.
They
a e
hus
o
less
bene i
o
g ey
long-ea ed
ba s
han
a e
me cu y
apou
ones.
The
e ec
o
s ee
lamps
on
P.
au i us
is
unce ain.
Rydell
and
Racey
(1995)
sugges ed
ha
some
species
may
a oid c ossing
li
a eas
and
may
hus
be
p e en ed
by
s ee
lamps
om
eaching
impo an
o aging
a eas.
This
has
no
been
shown
o
be
he
case
o
P.
au i-
us-,
Howa d
(1995)
ound
ha
indi iduals
commu ing
om
a
oos
o
o aging
a eas
we e
no
de e ed
by
a
powe ul
halogen
ligh
shining
ac oss
hei
no mal
lyway.
Howe e ,
i
is
possible
ha
a
s ee
lamp
may
a ac
mo hs
o
i
om
a
conside able
su ounding
a ea.
Da k
pa s
o
his
a ea
will
hus
con ain
ew
mo hs
and
so
o aging
he e
may
become
unp o i able.
Ba
Boxes
143
U baniza ion
P.
aus iacus
h i es
in
illages
and
small
owns
whe e
he e
a e
houses
in
which
o
oos
and
close-by
ga dens,
o cha ds
and
pa ks
in
which
o
o age.
I
also
appea s
o
be
well
adap ed
o
li ing
in
ci y
subu bs
in
some
egions
-
Gaisle
and
Baue o a
(1985-6)
ound
ou
nu s-
e y
colonies
in
buil -up
a eas
in
he
ci y
o
B no
and
equen ly
ne ed
indi iduals
in
he
subu bs.
Because
i
does
no
depend
on
woodland
bu
p e e s
o
o age
in
places
sue
as
ga dens
and
pa ks,
i
may
be
ound
in
mos
u ban
si ua ions
excep
inne
ci ies.
P.
au i us,
on
he
o he
hand,
depends
on
woodland
and
a ely
occu s
in
u ban
si ua-
ions.
Gaisle
and
Baue o a
(1985-6)
ound
hese
ba s
only
occasionally
a ound
B no
and
only
on
he
ex eme
ou e
limi s
o
he
ci y.
En wis le
e
al.
(1997)
ound
no
oos s
wi hin
he
ci y
o
Abe deen,
and
Hanak
(1969)
and
Gaisle
(1979)
simila ly
ound
no
oos s
wi hin
ci ies.
P.
au i us
h i es
in
u al
a eas
and
in
small
illages.
Inc easing
u baniza ion
in
many
coun ies
hus
p esen s
a
majo
h ea
o
he
species
and
has
been
sugges ed
as
a
eason
o
popula ion
declines
in
ecen
yea s
(Speakman
e
al.,
1991a).
BAT
BOXES
A i icial
oos s
in
he
o m
o
boxes
simila
o
bi d
nes ing
boxes
a e
used
by
ba s
in
some
a eas
and
schemes
in ol ing
hei
use
can
be
success ul
in
es ablishing
ba
popula ions,
pa -
icula ly
in
a eas
whe e
ew
o he
oos
si es
exis .
Ea ly
designs
o
ba
box
a e
simila
o
bi
boxes
excep
ha
en y
is
h ough
a
sli
a
he
bo om
ins ead
o
a
ound
hole
and
he
inside
o
he
box
is
g oo ed
o
allow
ba s
o
cling
o
i
(Figu e
9.2).
Boxes
a e
made
o
un ea ed
wood
and
hung
on
ees
o
buildings.
Thei
o ien a ion
does
no
seem
o
be
impo an ,
and
in
mos
schemes
boxes
a e
placed
in
g oups,
wi h
indi iduals
acing
in
se e al
di ec ions
so
ha
ba s
ha e
a
choice.
Boxes
ha e
a
emo able
lid
o
allow
inspec ion.
Thei
heigh
does
no
seem
o
be
c i ical
wi h
ega d
o
whe he
o
no
ba s
use
hem,
bu
in
gene al
hey
a e
placed
as
high
as
possible
in
ees
o
educe
he
dange s
o
p eda ion
o
andalism.
Apa
om
he
o iginal
wooden
boxes,
se e al
di e en
designs
a e
cu en ly
in
use,
including
one
made
o
a
mix u e
o
conc e e
and
sawdus
(Figu e
9.3),
which
appea s
o
be
success
u
in
a ac ing
ba s.
.
Ba
box
schemes
ha e
been
ound
o
be
mos
success ul,
in
e ms
o
ba s
using
hem,
when
hey
a e
es ablished
in
a eas
con aining
ew
buildings
which
could
be
used
as
oos s
and
no
ee
holes
o
o he
na u al
oos s.
La ge
coni e ous
plan a ions
a e
o en
in
isola ed
a eas
and
ee
holes
a e
a e
in
coni e s;
i
is
he e o e
no
su p ising
ha
he
mos
success-
ul
schemes
a e
in
coni e ous
o es s
(e.g.
Al ingham
and
Bullock,
1988;
Boyd
and
S ebbings,
1989;
Benzal,
1991).
Schemes
in
upland
a eas
ha e
been
epo ed
o
a ac
mo e
ba s
han
do
hose
a
lowe
al i ude
(Luge ,
1977;
Benzal,
1991),
and
his
has
been
a ibu ed
o
ewe
al e na i e
oos s
being
a ailable
a
highe
al i ude.
Apa
om
hei
conse a ion
pu pose
o
p o iding
oos s
and
a ac ing
ba s
o
new
a eas,
ba
boxes
a e
popula
wi h
bo h
he
public
and
ba
g oups
—
hey
a e
a
p ac ical
me hod
o
helping
in
ba
conse a ion
and,
when
hey
do
a ac
ba s,
hey
can
clea ly
be
seen
o
be
ha ing
a
posi i e
e ec .
They
can
also
p o ide
aluable
in o ma ion
on
ba
pop-
ula ions.
Long- e m
assessmen s
o
su i al,
sex
a ios
and
popula ion
dynamics
(Boyd
and
S ebbings,
1989;
Benzal,
1991)
ha e
been
made
using
ba
box
schemes.

144
Long-Ea ed
Ba s
and
Humans
10
cm
FIG
9.2
A
wooden
ba
box
wi h
one
side
emo ed,
showing
he
g oo ed
in e io ,
emo able
lid
and
en y
sli .
The
box
is
made
o
ough,
un ea ed
wood
and
is
nailed
o
a
ee
o
building.
Use
by
Long-ea ed
Ba s
P.
au i us
is
one
o
he
Eu opean
species
which
mos
equen ly
uses
ba
boxes
and
one
o
he
ew
which
o ms nu se y
colonies
in
hem.
Because
his
species
occupies
ee
holes,
ba
boxes
esemble
hei
na u al
oos s.
They
also
make
ex ensi e
use
o
woodland,
whe e
mos
ba
box
schemes
a e
si ua ed.
Al ingham
and
Bullock
(1988)
se
up
a
scheme
in
a
coni e -
ous
o es
in
sou h-eas
Sco land,
in
an
a ea
whe e
he e
we e
e y
ew
al e na i e
oos
si es.
Wi hin
3
yea s,
60%
o
he
boxes
had
been
used.
While
P.
pipis ellus,
he
mos
nume ous
species
in
he
boxes,
used
hem
only
o
ma ing,
he
wo
o he
species
ound,
P.
au i us
and
Myo is
na e e i,
bo h
o med
nu se y
colonies.
S ebbings
and
Walsh
(1983)
ound
ha
P.
au i us
was
he
commones
species
using
boxes
in
a
scheme
in
no he n
England,
while
Ba
Boxes
145
10
cm
FIG
9.3
A
conc e e-sawdus
ba
box,
consis ing
o
a
hollow
cylinde
wi h
wads
app oxima ely
4
cm
hick.
The e
is
a
sli
a
he
bo om
o
en y
and
he
oo
is
emo -
able
o
allow
inspec ion.
The
box
is
hung
om
a
ee
b anch
by
a
wi e
loop.
Bam o d
(1985)
epo ed
ha
his
species
was
ound
in
bi d
nes
boxes
in
a
la ch
(La ix
lep-
olepis)
plan a ion
in
Wales.
Schwenke
(1988)
es ima ed
ha
he
ins alla ion
o
ba
boxes
o
a
densi y
o
0.11
ha"'
and
he
inc ease
in
bi d
boxes
o
0.66
ha
in
a
o es
in
Ge many
inc eased
he
ba
popula ion
om
279
o
462
o e
a
pe iod
o
5
yea s.
In
ano he
pa
ol
Ge many,
Nagel
and
Nagel
(1988)
ound
ha
he
ins alla ion
o
ba
boxes
in
a
o es
a ea
d ama ically
inc eased
he
popula ion
o
ba s,
especially
o
P.
au i us.
Once
a ac ed
o e
a ea
by
hese
boxes,
ba s
also
oos ed
in
bi d
boxes
which
had
p e iously
been
p esen
bu
had
no
been
occupied
by
ba s.
Bo h
Schwenke
(1988)
and
Taake
and
Hildenhagen
(1989)
epo ed
ha
P.
au i us
oos ed p e e en ially
in
conc e e-sawdus
boxes
a he
han
in
wooden
ones.
I
aake
and
Hildenhagen
(1989)
sugges ed
his
may
ha e
been
because
conc e e-sawdus
boxes
ha e
146
Long-Ea ed
Ba s
and
Humans
be e
insula ion;
hey
hea up
mo e
slowly
han
wooden
ones
when
exposed
o
he
sun and
emain
wa m
o
longe ,
hus
unde going
less
luc ua ion
in
empe a u e.
P.
aus iacus
a ely
uses
ba
boxes.
As
a
s ic ly
house-dwelling
species
which
always
seeks
wa m
a ics
and
which
is
no
known
o
use
ee
holes,
i
appea s
o
a oid
boxes,
and
schemes
in
o es
a eas
ha e
had
no
success
in
a ac ing
i .
The
only
epo
o
g ey
long-
ea ed
ba s
being
ound
in
boxes
is
om
Poland
(Kowalski
and
Lesinski,
1994),
whe e
mo e
han
hal
he
21
species
known
o
occu
we e
ound
in
boxes,
a
leas
occasionally.
P.
aus-
iacus
used
hem
a ely,
and
we e
only
e e
ound
in
boxes
a ached
o
buildings.
Thus,
al hough
boxes
in
o es s
appea
o
be
o
no
use
o
conse a ion
o
his
species,
boxes
on
buildings
may
be
o
bene i
in
places
whe e
sui able
house
oos s
a e
in
sho
supply.
PRACTICAL
CONSERVATION
MEASURES
As
knowledge
o
he
biology,
ecology
and
beha iou
o
long-ea ed
ba s
accumula es,
i
is
becoming
easie
o
iden i y
he
mos
impo an
aspec s
o
hei
conse a ion
and
o
di ec
p ojec s
owa ds
hei
speci ic
needs.
While
ecen
legisla ion
has
concen a ed
on
summe
oos s,
i
is
now
becoming
ob ious
ha
o aging
habi a s
and
hibe nacula
a e
also
in
need
o
p o ec ion.
Fo aging
A eas
Deciduous
woodland
should
be
p o ec ed,
pa icula ly
a eas
close
o
P.
au i us
oos s.
Whe e
woodland
is
being
des oyed
on
a
la ge
scale,
i
is
some imes
possible
o
p ese e
pa s
o
i ,
and
e en
a
ai -sized
copse
ound
a
house
can
be
su icien
o
suppo
a
colony.
Hedges,
eelines
and
o he
ea u es
which
may
be
used
as
lyways
should
also
be
p ese ed,
since
he
absence
o
such
ea u es
can
make
pa ches
o
woodland
inaccessible.
Small-scale
p ojec s
o
plan
and
p o ec
joining
s ips
o
woodland
be ween
pa ches
o
o es
and
be ween
oos s
and
o aging
a eas
a e
cu en ly
being
ca ied
ou
in
many
a eas
and
a e
bene icial
o
long-ea ed
ba s.
Whe e
coni e ous
woodland
domina es,
he
quali y
can
be
imp o ed
by
di iding
dense
blocks
o
ees
up
by
clea ings
and
ides.
Such
p ac ices
will
inc ease
he
edge
habi a
a ail-
able
and
so
bene i
long-ea ed
ba s.
Ba
box
schemes
also
make
coni e ous
plan a ions
mo e
accessible
by
p o iding
oos
si es.
I
possible,
he
quali y
o
coni e ous
woodland
should
be
imp o ed
by
he
plan ing
o
some
deciduous
ees;
e en
a
line
ound
he
edge
is
o
bene i .
The
plan ing
o
mixed
o
o
deciduous
woodlands
in p e e ence
o
pu ely
coni e ous
ones
should
be
encou aged
whe e e
possible.
T ee
Holes
Hollow
ees
and
old,
deciduous
ees
wi h
hollow
b anches
should
be
p ese ed
whe e e
possible
as
po en ial
oos
si es.
Such
si es
a e
ypically
used
only
as
cool,
empo a y
oos s
(En wis le,
1994)
and
so
may
be
unoccupied
mos
o
he
ime;
his
does
no
make
hem
any
less
impo an
o
he
ba s.
Fuh mann
and
Godmann
(1991)
in es iga ed
di e en
ypes
o
ca i y
in
deciduous
ees
and
ound
ha
slow- lying
species
such
as
P.
au i us
and
Myo is
bechs einii
use
low
ca i ies,
such
as
a e
ound
in
old
coppiced
woods.
Old
ees
and
s umps
P ac ical
Conse a ion
Measu es
147
a e
he e o e
impo an
o
long-ea ed
ba s
and
should
be
p o ec ed.
When
hollow
ees
ha e
o
be
elled,
hey
should
always
be
ca e ully
checked
i s
o
oos ing
ba s
man)
unnecessa y
dea hs
occu
du ing
elling
ope a ions.
Roos s
Long-ea ed
ba s
show
a
high
deg ee
o
ideli y
o
nu se y
oos s.
They
ha e
been
shown
o
be
selec i e
in
picking
houses
in
which
o
oos
(En wis le
e ai,
1997)
and
sui able
houses
may
no
be
as
plen i ul
as
has
been
assumed
un il
ecen ly.
E ic ion
o
a
colony
could
he e-
o e
ha e
se ious
consequences
and
i
is
impo an
o
conse a ionis s
o
make
e e y
e o
o
pe suade
oos
owne s
o
be
ole an
o
hei
ba s.
Ad ice
and
suppo
a e
needed,
and
p ac ical
assis ance
such
as
wi h
emo ing
d oppings
om
a ics
can
make
he
di e ence
be ween
ba s
being
allowed
o
s ay
and
being
excluded.
Because
o
long-ea ed
ba s
ulne -
abili y
o
imbe
ea men
chemicals,
igilance
is
necessa y
in
oos s
o
ensu e
ha
ea -
men
is
ca ied
ou
wi h
app op ia e
chemicals
and
in
he
absence
o
ba s
P osecu ion
a e
ba s
a e
killed
may
p e en
u he
inciden s
bu
does
no
help
he
dea
a s.
e
ene gy
and
diligence
o
ba
g oup
membe s
is
in aluable
in
moni o ing
oos s
and
he
heal h
o
he
ba s
in
hem.
, ,
,
,
Old
a m
buildings,
like
ee holes,
a e
impo an
as
empo a y
oos s,
ihey
should
oe
p ese ed
whe e e
possible
and
checked
ca e iilly
o
signs
o
oos ing
ba s be o e
being
demolished
o
eno a ed.
Hibe nacula
Besides
p o ec ing
ca es
and
mines,
which
is
a
conse a ion measu e
needed
o
mos
hibe -
na ing
ba s,
help
can
also
be
p o ided
o
long-ea ed
ba s
by
building
a i icial
hibe nacula.
Se e al
p ojec s
in
Ge many
ha e
ecen ly
shown
he
bene i s
o
such
s uc u es.
e s
(1993,
1994)
eco ded
ha
a
conse a ion
g oup,
in
coope a ion
wi h
he
Ge man
a me
o ces,
eno a ed
se e al
old
mili a y
bunke s
and
disused
cella s
and
a
de elic
wa e
PU™P
ing
s a ion.
P.
au i us
was
he
species
which
made
mos
use
o
he
buildings,
an
a s
mo ed
in
du ing
he
i s
win e
a e
es o a ion
was
comple ed.
Use
was
ma
e
Y
a s
o
he
c e ices
inco po a ed
in o
he
ceilings
and
walls.
A
simila
p ojec
nea
o
ogne
(Buchen,
1992)
in ol ed
he
cons uc ion
o
an
a i icial
galle y
o
eplace
some
o
he
mines
and
qua ies
in
he
a ea
which
ha e
been
illed
in
o
blocked
up.
Two
yea s
a e
con-
s uc ion,
i e
species,
including
P.
au i us,
we e
ound
o
hibe na e
in
he
galle y.
O he
conse a ion
p ojec s
on
hibe nacula
ha e
included
su eys
o
amass
in
o ma ion
on
he
use
o
a ious
s uc u es.
Rydell's
(1989c)
su ey
o
s o age
cella s
in
Sweden
empha-
sized
hei
impo ance
o
P.
au i us
(Chap e
8)
and
Haensel
(1994)
epo ed
ha
se en
ba
species
hibe na ed
in
cella s
ound
B andenbu g
in
Ge many.
His
su ey
concen a e
on
collec ing
in o ma ion
which
could
be
used
o
es o e
cella s
in
such
a
way
as
o
make
em
sui able
as
ba
hibe nacula.
.
Ba
conse a ion
has
made
g ea
s ides
du ing
he
las
10
yea s.
App ecia ion
o
e
p oblem
o
declining
numbe s
o
indi iduals
and
he
need
o
ind
and
implemen
conse -
a ion
measu es
ha e
become
i mly
es ablished
and
he
public
a e
beginning
o
see
em
as
a
g oup
in
need
o
p o ec ion
and
no
as
a
pes .
Conside able
ad ances
a e
also
being
made
in
ou
unde s anding
o
he
ecological
equi emen s
o
ba s,
and
long-ea ed
ba s,
as
148
Long-Ea ed
Ba s
and
Humans
species
which
li e
in
close
p oximi y
o
humans,
a e
among
hose
whose
needs a e
begin-
ning
o
be
unde s ood.
They
a e
also
e y
a ac i e
animals
which
g ea ly
help
he
conse -
a ionis s'
a emp s
o
pe suade
he
public
o
become
mo e
'ba
iendly'.
The e
is,
howe e ,
no
oom
o
complacency
-
hese
ba s,
hei
oos s
and
hei
habi a s
a e
s ill
e y
much
in
need
o
ac i e
conse a ion.
Because
he
gene al
public
in e p e s
kindness
o
ani-
mals
as
meaning
kindness
o
nice
animals,
i
is
essen ial
o
po ay
long-ea ed
ba s
as
nice
animals.
I
is
he
esponsibili y
o
us
all
o
p o ec
and
conse e
ou
wildli e
and
o
ensu e
a
secu e
u u e
o
hese
beau i ul
and
ascina ing
ba s.
APPENDIX
1
Scien i ic
and
Common
Names
o
Ba
Species
An ozous
pallidas
Ba bas ella
ba bas ellus
Ca diode ma
co
Cloeo us
pe ci ali
Co yno hinus
mexicanus
Co yno hinus a inesquii
Co yno hinus
ownsendii
ingens
Co yno hinus
ownsendii ownsendii
Co yno hinus
ownsendii
i ginianus
Ep esicus
uscus
Ep esicus
nilssonii
Ep esicus
se o inus
Eude ma
macula um
Idionyc e is
phyllo us
Lasiu us
cine eus
Lasiu us
cine eus
semo us
La ia
ons
Miniop e us
sch eibe sii
Myo is
bechs einii
Myo is
bly hii
Myo is
dasycneme
Myo is
dauben onii
Myo is
ema gina us
Myo is
e o is
Myo is
luci ugus
Myo is
myo is
Myo is
mys acinus
Myo is
na e e i
Myo is
sep en ionalis
Myo is
sodalis
Nyc alus
noc ula
O omops
spp.
pallid
ba
ba bas elle
A ican
alse
ampi e
ba
sho -ea ed
iden
ba
Mexican
big-ea ed
ba
Ra inesque's,
o
eas e n,
big-ea ed
ba
Oza k
big-ea ed
ba
Townsend's,
o
wes e n,
big-ea ed
ba
Vi ginia
big-ea ed
ba
big
b own
ba
no he n
ba
se o ine
spo ed
ba
Allen's
big-ea ed
ba
hoa y
ba
Hawaiian
hoa y
ba
yellow-winged
ba
long- inge ed
ba
Bechs ein's
ba
lesse
mouse-ea ed
ba
pond
ba
Dauben on's
ba
no ch-ea ed
ba
long-ea ed
Myo is
li le
b own
ba
g ea e
mouse-ea ed
ba
whiske ed ba
Na e e 's
ba
no he n
long-ea ed
ba
social
ba
noc ule
big-ea ed
ee- ailed
ba s
149
150
Scien i ic
and
Common
Names
o iBa
Species
Pipis ellus
pipis ellus
Pleco us
au i us
Pleco us
aus iacus
Pleco us
ai anus
Pleco us
ene i ae
Rhinolophus
e umequinum
Rhinolophus
hipposide os
Tada ida
b asiliensis
Vespe ilio
mu inus
pipis elle
b own
long-ea ed
ba
g ey
long-ea ed
ba
Taiwan
long-ea ed
ba
Tene i e
long-ea ed
ba
g ea e
ho seshoe
ba
lesse
ho seshoe
ba
Mexican
ee- ailed
ba
pa icolou ed
ba
APPENDIX
2
Ca e
o
S anded
o
Inju ed
Ba s
STRANDED
BATS
O
CCASIONALLY,
ba s
en e
houses
o
o he
buildings
and
become
apped.
A e
ly-
ing
a ound
inside
o
some
ime,
hey
become
exhaus ed
and
may
la e
be
ound
and
escued.
Thei
immedia e
need
is
o
be
gi en
ood
and
wa e ,
a e
which
mos
eco e
apidly
and
can
be
eleased.
Wa e
is
bes
o e ed
on
a
e y
ine
pain b ush.
Ini ially,
his
can
be
b ushed
agains
he
ba 's
lips,
a e
which
he
ba
will
usually
lap
om
he
b ush.
In
summe ,
enough
insec s
o
e i e
a
ba
can
usually
be
caugh
wi h
a
bu e ly
ne .
Small
lies,
caddis
lies
o
mo hs
a e
all
accep ed
by
long-ea ed
ba s
and
a e
ela i ely
easy
o
collec ;
hey
a e
bes
o e ed
mashed
up
and
held
in
a
blun
pai
o
weeze s.
When
no
insec s
a e
a ail-
able,
mashed
mealwo ms
(see
below)
can
be
gi en
ins ead.
A
i s ,
ood
may
ha e
o
be
pushed
gen ly
in o
he
ba 's
mou h,
bu
as
i
eco e s
i
will
snap
a
i
a idly.
Hung y,
dehy-
d a ed
ba s a e
always
o pid
when
ound, bu
as
hey
a e
ed
hey
can
be
el
o
wa m
up.
Once
wa m
and
ully
ac i e,
hey
should
be
eleased
as
soon
as
possible.
I
he
whe eabou s
o
he
oos
is
known,
he
ba
can
be
placed
in
i .
I
no ,
i
should
be
eleased
a ound
dusk
ou side
he
building
whe e
i
was
ound.
P o ided
i
is
wa m
and
ac i e,
i
will
ly
om
an
ou s e ched
hand
o
i s own
acco d.
I
should
no
be
h own
in o
he
ai
because
i
i
is
no
ully
wa med
up
i
may
all
o
he
g ound
and
be
inju ed
o
e en
acciden ally
s ood
on.
INJURIES
Ba s
wi h
se ious
inju ies
o
in ec ions
equi e
a en ion
om
a
e e ina y
su geon.
Rela i ely
ew
e s
ha e
much
expe ience
in
dealing wi h
ba s,
bu
he e
a e
some
who
spe-
cialize
in
ea ing
his
g oup.
Local
RSPCA
o ice s,
he
Ba
Conse a ion
T us
(see
below)
o
local
e e ina y
su ge ies
can
ad ise
on
how
o ob ain
expe
ea men .
Wing
inju ies
a e
he
commones
cause
o
disablemen
in
ba s.
Those
wi h
b oken
inge
bones
usually
eco e
and
he
bones
some imes
heal
on
hei
own,
usually
h ough
he
o -
ma ion
o
callouses
ound
he
b eak.
In
such
cases,
he
ba
will
be
able
o
ly
again,
bu
many
ba s
wi h
wing
bone
ac u es
do
no
egain
he
abili y
o
ly.
B oken
hume i
o
o e-
a ms
a e
a
mo e
se ious.
These
can
some imes
be
pinned
in
la ge
species,
bu
in
ba s
as
small
as
Pleco us
pinning
is
a ely
success ul.
In
any
case,
in ec ion
is
always
a
se ious
isk,
and
o
many
ba s
wi h
hese
inju ies,
ampu a ion
o
he
wing
is
he
only
possible
ea -
men .
Many
wo ke s
eel
ha
humane
des uc ion
is
a
kinde
al e na i e
o
he
ba .
151
152
Ca e
o
S anded
o
Inju ed
Ba s
KEEPING
BATS
IN
CAPTIVITY
The
law
allows
sick
o
inju ed
ba s
o
be
kep
in
cap i i y
i
hey
do
no
ha e
a
ealis ic
chance
o
su i ing
in
he
wild,
bu
i
should
be
s essed
ha
hey
should
only
be
kep
cap-
i e
i
he e
eally
is
no
al e na i e.
Those
who
unde ake
he
ca e
should
be
awa e
ha
ba s
li e
long
li es
and
ha
he
commi men
could
be
o
a
numbe
o
yea s.
Many
ba
g oups
ha e
among
hei
membe s
an
expe
ca e
who
can
ad ise
on
keeping
ba s
and
who
may
be
p epa ed
o
ake
on
he
ca e
o
new
cases.
Cap i e
ba s
in
ba
g oups
ul il
a
use ul
public-
i y
unc ion
-
hey
can
be
aken
o
lec u es
and
demons a ions
o
enable
membe s
o
he
public
o
see
and
handle
li e
ba s.
Cages
made
o
wood
o
me al
lined
wi h
wood
a e
sui able
o
housing
long-ea ed
ba s.
The
wood
mus
be
un ea ed
and
ough
enough
o
allow
ba s
o
g ip.
Mesh
on
he
op
o
on
mus
be
ine
enough
o
p e en
hem
om
escaping
and
he e
should
be
a
da k
com-
pa men
o
sleeping.
Pape
owels
on
he
cage
loo
make
cleaning
easy.
Ba s
which
a e
capable
o
ligh
emain
much
heal hie
i
hey
ha e
daily
access
o
a
la ge
oom
o
enclo-
su e
whe e
hey
can
ly
eely
and
sa ely.
Wa e
mus
be
eely
a ailable,
and
is
bes
p o ided
in
a
shallow
dish
o
a
small
bi d
eede
a ached
o
he
side
o
he
cage.
Whe e
possible,
ba s
should
be
ed
on
eshly
caugh
insec s
such
as
mo hs
o
lies.
This
is
no
always
p ac icable
and
such
ood
can
be
supplemen ed
wi h
o
eplaced
by
mealwo ms
(Teneb io
spp.),
a ail-
able
om
pe
shops
o
by
mail
o de .
These
should,
howe e ,
be
supplemen ed
wi h
i a-
mins,
since
hey
lack
some
nu ien s
ound
in
wild
insec s.
Vi amin
supplemen s
such
as
Vi amen
may
be
sp inkled
on o
ood
o ,
in
soluble
o m,
added
o
d inking
wa e .
An
adul
P.
au i us
will
equi e
abou
25-30
mealwo ms
o
2.5-3.0
g
o
insec s
pe
day,
al hough
mos
ca e s
eed
o
appe i e,
pa icula ly
i
he
ba
is
able
o
ly
daily
-
hose
which
canno
ly
can
become
obese
i
o e ed.
REARING
BABY
BATS
Rea ing
abandoned
o
o phaned
in an s
is
ime-consuming,
di icul
and
equen ly
unsuc-
cess ul
-
again,
ba
g oups
may
be
able
o
p o ide
con ac
wi h
expe s
in
his
ield.
A
milk
subs i u e
has
o
be
adminis e ed
un il
he
in an
is
abou
4
weeks
old,
and
his
is
bes
gi en
h ough
a
pipe e
o
on
a
e y
ine
pain b ush.
The
mos
success ul
oods
a e
epo ed
o
be
dilu ed
cow's
milk
(1:1
milk
:
wa e )
o
a
bi ch-milk
subs i u e
such
as
'Lac ol'.
Baby
ba s
ha e
o
be
ed
a
leas
e e y
2-3
h
h oughou
he
day
and
a e
eeding he
baby
should
be
washed
e y
gen ly
wi h
co on
buds
dipped
in
wa e
o
simula e
he
g ooming
ac ion
o
i s
mo he .
In an s
should
be
kep
a
a
cons an
empe a u e
o
abou
30°C.
Weaning
is
also
a
di icul
p ocess,
since
many
cap i e
ju eniles
seem
e y
eluc an
o
s a
ea ing
mealwo ms
o
insec s
-
much
ime
and
pa ience
is
needed.
I
is
gene ally
conside ed
ha
ba s
ea ed
in
cap i i y
should
no
be
eleased;
ba s
ha e
complex
social
beha iou ,
much
o
which
is
lea ned
om
conspeci ics,
and
hand- ea ed
in an s
do
no
acqui e
his
lea ning
and
a e
unlikely
o
su i e
in
he
wild.
APPENDIX
3
Use ul
Con ac
Add esses
in
he
UK
The
Ba
Conse a ion
T us
(BCT)
15
Clois e s
House,
8
Ba e sea
Pa k
Road,
London
SW8
4BG.
Telephone
0171
627
2629.
Fax
0171
627
2628.
BCT
is
in ol ed
in
educa ion,
ad ice
and
p omo ion
o
conse a ion
o
ba s
in
B i ain.
I
ac s
as
a
coo dina ion
cen e
o
ba
g oups
and
is
also ac i ely
in ol ed
in
esea ch
conce ned
wi h
ba
conse a ion.
Local
Ba
G oups
The e
a e
a ound
90
o
hese,
co e ing
mos
a eas
in
he
coun y.
The
names
and
add esses
o
con ac
pe sons
o
indi idual
g oups
can
be
ob ained
om
BCT.
The
Mammal
Socie y
15
Clois e s
Business
Cen e,
8
Ba e sea
Pa k
Road,
London
SW8
4BG.
Telephone
0171
498
4358.
Fax
0171
498
4459.
This
socie y
p omo es
he
s udy
o
mammals,
including
ba s,
by
ama eu
na u alis s
and
p o essional
scien is s.
I
designs
and
supplies
ba
ings
and
also
p oduces
a
numbe
o
publica ions
ele an
o
he
s udy
o
ba s.
Sco ish
Na u al
He i age
(SNH)
12
Hope
Te ace,
Edinbu gh
EH9
2AS.
Telephone
0131
554
9797.
English
Na u e
No hmins e
House,
Pe e bo ough
PE1
1UA.
Telephone
01733
340345.
Coun yside
Council
o
Wales
(CCW)
Plas
Pen hos,
F o dd
Pen hos,
Bango ,
Gwynedd
LL57
2LQ.
Telephone
01248
370444.
These
h ee
s a u o y
na u e
conse a ion
o ganiza ions
a e
esponsible
o
ad ising
Go e nmen
on
na u e
conse a ion
in
he
UK.
They
gi e
ad ice
on
ba
p oblems
unde
he
Wildli e
and
Coun yside
Ac
1981,
en o ce
p o ec ion
o
all
ba s
unde
his
law
and
also
adminis e
he
licensing
sys em.
Wo ld
Wide
Fund
o
Na u e
(WWF)
Panda
House,
Weyside
Pa k,
Godalming,
Su ey
GU7
1XR.
Telephone
01483
426444.
Fauna
&
Flo a
In e na ional
(FFI)
G ea
Eas e n
House,
Tenison
Road,
Camb idge
CB1
2DT.
Telephone
01223
571000.
Vincen
Wildli e
T us
(VWT)
10
Lo a
Lane,
London
EC3R
8DT.
Telephone
0171
283
2089.
The
abo e
h ee
olun a y o ganiza ions
all
include
p omo ion
o
ba
conse a ion
in
hei
wo k.
153

154
Use ul
Con ac
Add esses
in
he
UK
Royal
Socie y
o
he
P e en ion
o
C uel y
o
Animals
(RSPCA)
Causeway,
Ho sham,
Sussex
RH12
1HG.
Telephone
01403
264181.
Sco ish
Socie y
o
he
P e en ion
o
C uel y
o
Animals
(SSPCA)
19
Mel ille
S ee ,
Edinbu gh
EH3
7PL.
Telephone
0131
225
6418.
Re e ences
Abelence ,
V.,
Pidoplicko,
I.G.
and
Popo ,
B.M.
(1956).
Fauna
Uk ainy
1.
Ssa ei.
Insec i o a,
Chi op e a.
AN.
USSR
Ins .
Zooh
229—446.
Achy a,
L.
and
Fen on,
M.B.
(1992).
Echoloca ion
beha iou
o
espe ilionid
ba s
(.Lasiu us
cine eus
and
Lasiu us
bo ealis)
a acking
ai bo ne
a ge s
including
a c iid
mo hs.
Can.
J.
Zool.,70:1292-1298.
Adam,
M.D.,
Lacki,
M.J.
and
Ba nes,
T.G.
(1994).
Fo aging
a eas
and
habi a
use
o
he
Vi ginia
big-ea ed
ba
in
Ken ucky./.
Wildli e
Managemen ,
58(3):
462-469.
Aellen,
V.
(1971).
Le
chau e-sou is
Pleco us
aus iacus
(Fische )
en
Suisse.
Cong .
Suisse
SpeUol.
Neucha el,
1970,
Ac .4:
167-172.
Aellen,
V.
(1983-4).
Mig a ions
de
chau es-sou is
en
Suisse.
No e
complemen ai e.
Myo is,
21-22:
185-189.
Agee,
H.R.
(1967).
Response
o
he
acous ic
cell
o
he
bollwo m
and
obacco
budwo m
o
ul asound./.
EconomicEn omol.,
60:
366-369.
Agee,
H.R.
(1969).
Response
o
lying
bollwo m
mo hs
and
o he
ympana e
mo hs
o
pulsed
ul asound.
Ann.
En omol.
Soc.
Ame ica,
62:
801—807.
Ahlen,
I.
(1981).
Iden i ica ion
o
Scandina ian
ba s
by
hei
sounds.
Repo
No.6,
Dep .
Wildli e
Ecol.
Swedish
Uni .
Ag ic.
Sci.
Ahlen,
I.
and
Ge ell,
R.
(1989).
Dis ibu ion
and
s a us
o
ba s
in
Sweden.
In:
Eu opean
Ba
Resea ch
1
W(Hanak,
V.,
Ho acek,
I.
and
Gaisle ,
J.,
eds).
Cha les
Uni e si y
P ess,
P ague:
319-325.
Albe s,
S.
(1993)
Bau
on
Flede mauswin e qua e ie en
du ch
die
Bundesweh
in
Muns e
(Landk eis
Sol au
Fallingbos el).
Nyc alus,
Be lin,
4:
462—464.
Albe s,
S.
(1994).
Bau
on
un
Flede maus-Win e qua ie en
in
Raum
Bispingen,
Landk eis
Sol au-Fallingbos el
(no dos liches Niede sachensen).
Nyc alus,
Be lin,
5:
191-195.
Ald idge,
H.D.J.N.
(1987).
Tu ning
ligh
o
ba s./.
Exp.
Biol.,
128:
419^25.
Ald idge,
H.D.J.N.
and
B igham,
R.M.
(1988).
Load
ca ying
and
manoeu e abili y
in
an
insec i o ous
ba :
a
es
o
he
5%
' ule'
o
adio- eleme y./.
Mamm.,
69:
378.
Al ingham,
J.D.
(1996).
Ba s.
Biology
and
Beha iou .
Ox o d
Uni e si y
P ess,
Ox o d.
Al ingham,
J.
and
Bullock,
D.
(1988).
Ba
boxing
in
Fi e.
Ba cha
(NCC),
11:
4-7.
Ande ka,
F.W.
and
Angeh n,
P.
(1992).
T ansmi e
a achmen
me hods.
In:
Wildli e
Teleme y.
Remo e
Moni o ing
and
T acking
o
Animals
(P iede,
I.G.
and
Swi ,
S.M.,
eds).
Ellis
Ho wood,
Chiches e :
135-146.
Ande son,
M.E.
and
Racey,
P.A.
(1991).
Feeding
beha iou
o
cap i e
b own
long-ea ed
ba s,
Pleco us
au i us.
Anim.
Beha .,
42:
489—493.
Ande son,
M.E.
and
Racey,
P.A.
(1993).
Disc imina ion
be ween
lu e ing
and
non- lu e ing
mo hs
by
b own
long-ea ed
ba s,
Pleco us
au i us.
Anim.
Beha .,
46:
1151-1155-
155
156
Re e ences
An hony,
E.L.P.
(1987).
The
ole
o
he
an e io
pi ui a y
and
he
hypo halamus
in
con olling
ep oduc i e
cycles
in
ba s.
In:
Recen
Ad ances
in
he
S udy
o
Ba s
(Fen on,
M.B.,
Racey,
P.A.
and
Rayne ,
J.M.V.,
eds).
Camb idge
Uni e si y
P ess,
Camb idge:
421-439.
A le az,
R.
(1996a).
Feeding
beha iou
and
o aging
s a egy
o
ee-li ing
mouse-ea ed
ba s,
Myo is
myo is
and
Myo is
bly hii.
Anim.
Beha .,
51:
1-11.
A le az,
R.
(1996b).
Fo aging
beha iou
o
he
gleaning
ba
Myo is
na e e i
(Chi op e a,
Vespe ilionidae)
in
he
Swiss
Alps.
Mammalia,
60(2):
181-186.
A le az,
R.
and
Pe in,
N.
(1995).
The
ophic
niches
o
sympa ic
sibling
Myo is
myo is
and
M.
bly hii:
do
mouse-ea ed
ba s
selec
p ey?
In:
Ecology,
E olu ion
and
Beha iou
o
Ba s
(Racey,
P.A.
and
Swi ,
S.M.,
eds).
Symp.
Zool.
Soc.
London,
67:
361-376.
A nold,
H.R.
(1993).
A las
o
Mammals
in
B i ain.
Ins .
Te .
Ecol.,
London:
64—67.
Ascho ,
J.
(1966).
Ci cadian
ac i i y
pa e n
wi h
wo
peaks.
Ecology,
47(4):
657-662.
Aude ,
D.
(1990).
Fo aging
beha iou
and
habi a
use
by
a
gleaning
ba ,
Myo is
myo is
(Chi op e a,
Vespe ilionidae)./
Mamm.,
71(3):
420-427.
A e y,
M.I.
(1985).
Win e
ac i i y
o
pipis elle
ba s./.
Anim.
Ecol.,
54:
721-738.
Azza oli-Pucce i,
M.L.
and
Za a,
B.
(1988).
New
da a
on
he
Chi op e a
o
he
Cape
Ve de
Islands
(No h
A lan ic).
Mus.
Reg.
di
Scienze
Na .
Boll.
(To ino),
6(2):
603-616.
Baagoe,
H.J.
(1980-81).
Danish
ba s
,
s a us
and
p o ec ion.
Myo is,
18-19:
16-17.
Baagoe,
H.J.
(1987).
The
Scandina ian
ba
auna:
adap i e
wing
mo phology
and
ee
ligh
in
he
ield.
In:
Recen
Ad ances
in
he
S udy
o
Ba s
(Fen on,
M.B.,
Racey,
P.A.
and
Rayne ,
J.M.V.,
eds).
Camb idge
Uni e si y
P ess,
Camb idge:
57-74.
Bagley,
F.
and
Jacobs,
J.
(1985).
Census
echnique
o
endange ed
big-ea ed
ba s
p o ing
suc-
cess ul.
Endange ed
Species
Techn.
Bull.,
10(3):
5-7.
Bake ,
J.M.
and
Be y,
R.W.
(1980).
Syn he ic
py e h oid
insec icides
as
eplacemen s
o
chlo-
ina ed
hyd oca bons
o
he
con ol
o
wood-bo ing
insec s.
Holz
als
Roh-
und
We ks oj ,
38:
121-127.
Bake ,
R.J.
(1970).
Ka yo ypic
ends
in
ba s.
In:
Biology
o
Ba s,
Vol.
1
(Wimsa ,
W.A.,
ed.).
Academic
P ess,
New
Yo k:
65-96.
Bam o d,
R.
(1985).
Nes boxes
in
o es y
plan a ions.
Qua .
J.
Fo es y,
79(3):
153-158.
Ba a aud,
M. (1990).
Elemen s
su
le
compo emen
alimen ai e
des
O eilla ds
b un
e
g is,
Pleco us
au i us
(Linnaeus,
1758)
e
Pleco us
aus iacus
(Fische ,
1829).
Le
Rhinolophe,
7:
3-10.
Ba bou ,
R.W.
and
Da is,
W.H.
(1969).
Ba s
o
Ame ica.
Uni e si y
P ess,
Ken ucky,
Lexing on.
Ba bou ,
R.W.,
Da is,
W.H.
and
Hassell,
M.D.
(1966).
The
need
o
ision
in
homing
by
Myo is
sodalis.
J.
Mamm.,
47(2):
356-357.
Ba clay,
R.M.R.
(1991).
Popula ion
s uc u e
o
empe a e
zone
insec i o ous
ba s
in
ela ion
o
o aging
beha iou
and
ene gy
demand./.
Anim.
Ecol.,
60:
165-178.
Ba clay,
R.M.R.
(1995).
Does
ene gy
o
calcium
a ailabili y
cons ain
ep oduc ion
in
ba s?
In:
Ecology,
E olu ion
and
Beha iou
o
Ba s
(Racey,
P.A.
and
Swi ,
S.M.,
eds).
Symp.
Zool.
Soc.
London,
67:
245-258.
Ba clay,
R.M.R.
and
Bell,
G.P.
(1988).
Ma king
and
obse a ional
echniques.
In:
Ecological
and
Beha iou al
Me hods
o
he
S udy
o
Ba s
(Kunz,
T.H.,
ed.).
Smi hsonian
Ins i u e
P ess,
Washing on:
59-76.
Ba e -Hamil on,
G.E.H.
(1907).
Desc ip ion
o
wo
new
species
o
Pleco us.
Ann.
&
Mag.
Na . His ., 20:
520-521.
Ba e -Hamil on,
G.E.H.
(1910).
A
His o y
o
B i ish
Mammals.
1.
Ba s.
Gu ney
&
Jackson,
London.
Re e ences
157
Baue ,
K.
(I960).
Die
Sauge ie e
des
Neusiedle see-Gebie es
(Os e eich).
Bonn.
Zool.
Bei .,
11:
141-344.
Baue o a,
Z.
(1982).
Con ibu ion
o
he
ophic
ecology
o
he
g ey
long-ea ed
ba ,
Pleco us
aus iacus.
Fol.
Zool.,
31(2):
113-122.
Baue o a,
Z.
and
Zima,
J.
(1988).
Seasonal
changes
in
isi s
o
a
ca e
by
ba s.
Fol.
Zool.,
37(2):
97-111.
Beck,
A.
(1995).
Faecal
analysis
o Eu opean
ba
species.
Myo is,
32-33:
109-119.
Bekke ,
J.P.
and
Mos e ,
K.
(1990).
P eda ion
on
ba s
in
he
Ne he lands:
ac s
and
assump-
ions.
Abs ac ,
5 h
Eu opean
Ba
Res.
Symp.
Nybo g,
Denma k.
Bell,
T.
(1874).
A
His o y
o
B i ish
Quad upeds.
Van
Voo s ,
London.
Benzal,
J.
(1991).
Popula ion
dynamics
o
he
b own
long-ea ed
ba
{Pleco us
au i us)
occupy-
ing
bi d
boxes
in
a
pine
o es
plan a ion
in
cen al
Spain.
Ne h.
J.
Zool.,
41(4):
241-249.
Be na d,
R.,
Glazaczow,
A.
and
Samolag,
J.
(1991).
The
o e win e ing
ba
colony
in
S zaliny
(no hwes e n
Poland).
Ac a.
Zool.
C aco iensa,
34(2):
453-461.
Be y,
R.W.
(1983).
Recen
de elopmen s
in
he
emedial
ea men
o
wood-bo ing
insec
in es a ions.
Biode e mina ion,
5:
154—165-
Be and,
A.
(1992).
Donnees
p eliminai es
su
les
chau es-sou is
de
la
hau e
chain
py eneenne
(Py enees
a giegeoises).
Mammalia,
56(2):
290—292.
Bes ,
T.L.,
Ca ey,
S.D.,
Caesa ,
K.G.
and
Hen y,
T.H.
(1992).
Dis ibu ion
o
ba s
(Mammalia:
Chi op e a)
in
coas al
plain
ca es
o
sou he n
Alabama.
Na .
Spel.
Soc.
Bull.,
54(2):
61-65.
BezemJ.J.,
Slui e ,
J.W.
and
an
Hee d ,
P.F.
(1964).
Some
cha ac e is ics
o
he
hibe na ing
loca ions
o
a ious
species
o
ba s
in
Sou h
Limbu g.
1.
P oc.
Koninkl.
Akademie
an
We enschappen
Ams e dam
C,
67(5):
325—350.
Billing on,
G.
(1993).
BAT
g oups,
No.7.
Ba
Conse a ion
T us ,
London.
Bingley,
W.
(1809).
Memoi s
o
B i ish
Quad upeds.
Da on
&
Ha ey,
London.
Blake,
D„
Hu son,
A.M.,
Racey,
P.A.,
Rydell,
J.
and
Speakman,
J.R.
(1994).
Use
o
lampli
oads
by
o aging
ba s
in
sou he n
England.
J.
Zool.
London,
234:
453-^462.
Bogdanowicz,
W.
(1983).
Communi y
s uc u e
and
in e speci ic
in e ac ions
in
ba s
hibe na -
ing
in
Poland.
Ac a.
The iol.,
28:
357—370.
Bogdanowicz,
W.
and
Owen,
R.D.
(1996).
Landma k-based
size
and
shape
analysis
in
sys em-
a ics
o
he
pleco ine
ba s.
In:
Ad ances
in
Mo phome ies
(Ma cus,
L.F.
e
al.,
eds).
Plenum
P ess,
New
Yo k.
Bogdanowicz,
W.
and
U banczyk,
Z.
(1983).
Some
ecological
aspec s
o
ba s
hibe na ing
in
ci y
o
Poznum.
Ac a.
The iol.,
28:
371—385-
Boo h,
C.
and
Boo h,
J.
(1994).
The
Mammals
o
O kney.
Ki kwall.
Bo g,
J.,
Fio e,
M.,
Violani,
C.
and
Za a,
B.
(1990).
Obse a ions
on
he
Chi op e o auna
o
Gozo,
Mal ese
Islands.
Mus.
Reg
Sci.
Na .
Boll.
(To ino),
8(2):
501—516.
Boyd,
I.L.,
Myhill,
D.G.
and
Mi chell-Jones,
A.J.
(1988).
Up ake
o
Gamma-HCH
by
pip-
is elle
ba s
and
i s
e ec
on
su i al.
En .
Pollu ion,
51:
95-111.
Boyd,
I.L.
and
S ebbings,
R.E.
(1989).
Popula ion
changes
o
b own
long-ea ed
ba s
{Pleco us
au i us)
in
ba
boxes
in
The o d
Fo es ./.
Appl.
Ecol.,
26:
101-112.
Boye,
P.,
Po -Do e ,
B.,
Do e ,
K.
and
Deme opoulous,
A.
(1990).
New
eco ds
o
ba s
(Chi op e a)
om
Cyp us
and
no es
on
hei
biology.
Myo is,
28:
93-100.
B ack,
V.
and
Twen e,
J.W.
(1985).
The
du a ion
o
he
pe iod
o
hibe na ion
o
h ee
species
o
espe ilionid
ba s.
1.
Field
s udies.
Can.
J.
Zool.,
63:
2952—2954.
B adbu y,
J.W.
(1977a).
Lek
ma ing
beha iou
in
he
hamme -headed
ba .
Z.
Tie psychol.,
45:
225-255.
158
Re e ences
B adbu y,
J.W.
(1977b).
Social
o ganiza ion
and
communica ion.
In:
Biology
o
Ba s,
Volume
3
(Wimsa ,
W.A.,
ed.).
Academic
P ess,
New
Yo k:
1-72.
B igham,
R.M.
(1987).
The
signi icance
o
win e
ac i i y
by
he
big
b own
ba
{Ep esicus
Jus-
cus):
he
in luence
o
ene gy
ese es.
Can.
J.
Zool.,
65:
1240-1242.
B osse ,
A.
(1966).
La
Biologie
des
Chi op e es.
Masson,
Pa is:
240.
B own,
J.,
Flinde s,
I.
and
Richa dson,
P.W.
(1983).
The
use
o
chu ch
po ches
by
ba s./.
Zool.
London,
200:
292-295.
B own,
L.N.
and
B own,
C.K.
(1993).
Fi s
eco d
o
he
eas e n
big-ea ed
ba
(Pleco us
a inesquii)
in
sou he n
Flo ida.
Flo ida
Scien is ,
56(1):
63—64.
B own,
P.
(1976).
Vocal
communica ion
in
he
pallid
ba ,
An ozouspallidus.
Z.
Tie psychol.,
41:
34-54.
B own,
P.E.
and
G innell,
A.D.
(1980).
Echoloca ion
on ogeny
in
ba s.
In:
Animal
Sona
Sys ems
(Busnel,
R.G. and
Fish,
J.F.,
eds).
Plenum
P ess,
New
Yo k:
355-377.
B uijn,
Z.
(1990).
Domes ic
ca
Felis
ca usas
a
p eda o
o
ba s.
Lu a,
33(1):
30-34.
Buchen,
C.
(1992).
Bau
eines
kuns lichen
S ollens
als
Flede maus-Win e qua ie .
Nyc alus,
Be lin,
4:
269-273-
Buchle ,
E.R.
(1976).
A
chemiluminescen
ag
o
acking
ba s
and
o he
small
noc u nal
ani-
mals./.
Mamm.,
57:
173-176.
Bullock,
D.J.,
Combes,
B.A.,
Eales,
L.A.
and
P i cha d,
J.S.
(1987).
Analysis
o
he
iming
and
pa e n
o
eme gence
o
he
pipis elle
ba
(Pipis ellus
pipis ellus).
J.
Zool.
London,
211:
267-274.
Cas o ,
C.,
De me ,
K.
and
Jup ne ,
S.
(1993).
Von Tagesmenu
zum
Ges amp abspec um
des
G auen
Langoh s
{Pleco us
aus iacus)
-
2
Jah e
F eilanda bei
u den
Flede mausschu z.
Nyc alus,
Be lin,
4:
495-538.
Ca o,
C.
(1994).
Ba
De ec o
Manual.
Ba
Conse a ion
T us ,
London.
Ca o,
C.M.C.,
Racey,
P.A.
and
S ephenson,
P.J.
(1995).
Ac i i y
pa e ns
o
he
se o ine
ba
{Ep esicus
se o inus)
a
a
oos
in
sou he n
England./.
Zool.
London,
235:
635-644.
Cla k,
B.S.,
Leslie,
D.M.
and Ca e ,
T.S.
(1993).
Fo aging
ac i i y
o
adul
emale
Oza k
big-
ea ed
ba s
{Pleco us
ownsendii
ingens)
in
summe ./.
Mamm.,
74(2):
422—427.
Clee es,
T.R.
(1969).
He ing
gull
ca ching
and ea ing
ba .
B i ish
Bi ds,
62:
333.
Cock um,
E.L.
(1956).
Homing,
mo emen s
and
longe i y
o
ba s./.
Mamm.,
37(1):
48-57.
Coles,
R.B.,
Guppy,
A.,
Ande son,
M.E.
and
Schlegel,
P.
(1989).
F equency
sensi i i y
and
di ec ional
hea ing
in
he
gleaning
ba
Pleco us
au i us
(Linnaeus
1758)./.
Comp.
Physiol.
A,
165:
269-280.
Co be ,
G.B.
(1964).
The
g ey
long-ea ed
ba
Pleco us
aus iacus
in
England
and
he
Channel
Islands.
P oc.
Zool.
Soc.
London,
143(3):
511-515.
Co be ,
G.B.
(1970).
Vag an
ba s
in
She land
and
he
No h
Sea.
/
Zool.
London,
161:
281-282.
Co be ,
G.B.
(1971).
P o isional
dis ibu ion
maps
o
B i ish
mammals.
Mamm.
Re .,
1:
95-142.
Co be ,
G.B.
and
Hill,
F.E.
(1991).
A
Wo ld
Lis
o
Mammalian
Species,
3 d
edn.
B i ish
Museum
(Na u al
His o y),
London.
Co mack,
R.M.
(1964).
Es ima es
o
su i al
om
he
sigh ing
o
ma ked
animals.
Biome ika,
51:429-434.
C anb ook,
Ea l
o
(1963a).
Folding
o
he
ea s
o
a
long-ea ed
ba
{Pleco us
au i us).
Su olk
Na .
T ans.,
11(5):
386-387.
C anb ook,
Ea l
o
(1963b).
No es
on
he
eeding
habi s
o
he
long-ea ed
ba .
Su olk
Na .
T ans.,
11(2):
1-3.
Re e ences
159
C uci i,
P.
(1989).
Dis ibu ion,
di e si y
and
abundance
o
ca e
ba s
in
La ium
^ ai^-
In:
Eu opean
Ba
Resea ch
1987
(Hanak,
V.,
Ho acek,
I.
and
Gaisle ,
J.,
eds).
Cha les
Da^jT(1970).
Pho og aphic
eco ding
o
na u al
ac i i y
in
hibe na ing
ba s.
Bijd.
Die k.,
40:
Daan~S6
(1973).
Ac i i y
du ing
na u al
hibe na ion
in
h ee
species
o
espe ilionid
ba s.
DaanTs.
{l980)'
Long
J m
changes
in
ba
popula ions
in
he
Ne he lands:
a
summa y.
Lu a,
Daan,
S.^nd
Wiche s,
H.J.
(1968).
Habi a
selec ion
o
ba s
hibe na ing
in
a
limes one
ca e.
Sonde d uck
aus
Z.
Sauge ie kunde
Bd.,
33(5):
262-287.
u-
l
-c ,,
n
Dapson,
R.W.,
S udie ,
E.H.,
Buckingham,
M.J.
and
S ud.e ,
A.L.
(1977).
Hls °^em^ >
odi e ous
sec e ions
om
he
in egumen a y
glands
in
h ee
species
o
ba s./.
Mamm.,
58.
Da is,
R5
(1966).
Homing
pe o mance
and
homing
abili y
in
ba s.
Ecol.
Monog aphs,
36(3):
2Q2
237
Da is,
R.B.,
He eid,
C.F.
and
Sho ,
H.L.
(1962).
Mexican
ee- ailed
ba s
in
Texas.
Ecol.
Demlse^AT.
a980).
The
ba s
o
I an:
sys ema ics,
dis ibu ion,
ecology.
Fieldiana
Zool.
N.S.
De
Comsey,
G.
and
de
Cou sey,
P.J.
(1964).
Adap i e
aspec s
o
ac i i y
hy hms
in
ba s.
Biol.
Bull.,
126:
14-27.
,
.
,
. .
,
.
,
De
Fanis,
E.
and
Jones,
G.
(1995a).
Pos -na al
g ow h,
mo he -in an
in e ac ions
and^d
opmen
o
ocaliza ions
in
he
espe ilionid
ba
Pleco us
au i us.
J.
Zool.
London,
235-
83
)/.
De
Fanis,
E.
and
Jones,
G.
(1995b).
The
ole
o
odou
in
he
disc imina ion
o
conspecihcs
by
pipis elle
ba s.
Anim.
Beha .,
49:
835-839.
De
Paz,
O.
(1984).
On
he
dis ibu ion
o
genus
Pleco us
(Chi op e a:
Vespe ihomdae)
in
he
Ibe ian
Peninsula
and
Balea ic
Isles.
Mammalia,
48:
585-591.
Dulic
B
(1980).
Mo phological
cha ac e is ics
and
dis ibu ion
o
Pleco us
au i us
and
Pleco us
aus iacus
in
some
egions
o
Yugosla ia.
P oc.
5 h
In .
Ba
Res.
Con e ence-151-181.
Dunning,
D.C.
and
Roede ,
K.D.
(1965).
Mo h
sounds
and
he
insec -ca ching
beha iou
ol
ba s.
Science,
New
Yo k,
147:
173—174.
Eisen au ,
M.
(1936).
Bei ag
zu
Mechanik
des
Flede maus luges.
Z
wiss.
Zool.,
139—188
Eisen au ,
M.
(1937).
Die
Wi kung
neid ige
Tempe a u en
au
die
Emb yonen wichlung
bei
Flede mausen.
Biol.
Zbl.,
57:
59—74.
Eisen au ,
M.
(1950).
Die
E nahung
de
Flede mause.
Zool.
Jb.,
79:
115-177.
Ekman,
M.
and
de
Jong,
J.
(1996).
Local
pa e ns
o
dis ibu ion
and
esou ce
u iliza ion
o
lou
ba
species
{Myo is
b and i,
Ep esicus
nilssoni,
Pleco us
au i us
and
Pipis ellus
pipis ellus)
in
pa chy
and
con inuous
en i onmen s./.
Zool.
London,
238:
571-580.
Ellis,
S.E.
(1993).
Tabanidae
as
die a y
i ems
o
Ra inesque's
big-ea ed
ba :
implica ions
lo
i s
o aging
beha iou .
En omol.
News,
104(3):
118-122.
.
•
l
En wis le,
A.C.
(1994).
Roos
ecology
o
he
b own
long-ea ed
ba
{Pleco us
au i us)
in
no h-
eas
Sco land.
Unpublished
PhD
hesis,
Uni e si y
o Abe deen,
UK.
En wis le,
A.C.,
Racey,
P.A.
and
Speakman,
J.R.
(1996).
Habi a
exploi a ion
by
a
gleaning
ba ,
Pleco us
au i us.
Phil.
T ans.
R.
Soc.
Lond.
B,
351:
921—931.
En wis le,
A.C.,
Racey,
P.A.
and
Speakman,
J.R.
(1997).
Roos
selec ion
by
he
b own
long-
ea ed
ba
{Pleco us
au i us).
J.
Appl.
Ecol.,
34:
399-408.
160
Re e ences
En wis le,
A.C.,
Racey,
P.A.
and
Speakman,
J.R.
(in
p ess).
The
ep oduc i e
cycle
and
de e -
mina ion
o
sexual
ma u i y
in
male
b own
long-ea ed
ba s,
Pleco us
au i us.
J.
Zool.
London.
En wis le,
A.C.,
Speakman,
J.R.
and
Racey,
P.A.
(1994).
E ec
o
using
he
doubly
labelled
wa e
echnique
on
long- e m
ecap u e
in
he
b own
long-ea ed
ba
(Pleco us
au i us).
Can.
J.
Zool.,
72:
783-785.
E ke ,
H.G.
(1982).
Ecological
aspec s
o
ba
ac i i y
hy hms.
In:
Ecology
o
Ba s
(Kunz,
T.H.,
ed).
Plenum
P ess,
New
Yo k:
201-242.
E ke ,
H.G.
and
K ach ,
S.
(1978).
E idence
o
ecological
adap a ion
o
ci cadian
sys ems.
Oecologia
(Be l),
32:
71-78.
Fa ney,
J.
and
Fleha y,
E.D.
(1969).
Aspec
a io,
loading,
wing
span
and
memb ane
a eas
o
ba s.
/
Mamm.,
50(2):
362-367.
Fau e,
P.A.,
Fulla d,
J.H.
and
Dawson,
J.W.
(1993).
The
gleaning
a acks
o
he
no he n
long-
ea ed
ba ,
Myo issep en ionalis,
a e
ela i ely
inaudible
o
mo hs./.
Exp.
Biol.,
178:
173-189.
Fedyk,
A.
and
Fedyk,
S.
(1970).
Ka yo ypes
o
some
species
o
espe ilionid
ba s
om
Poland.
Ac a
The iol.,
15:
295-305.
Fedyk,
S.
and
Rup ech ,
A.L.
(1983).
Ch omosomes
o
some
species
o
espe ilionid
ba s.
2.
E olu iona y
ela ionships
o
pleco ine
ba s.
Ac a
The iol.,
28(10):
171-182.
Fen on,
M.B.
(1995).
Cons ain
and
lexibili y
—
ba s
as
p eda o s,
ba s
as
p ey.
In:
Ecology,
E olu ion
and
Beha iou
o
Ba s
(Racey,
P.A.
and
Swi ,
S.M.,
eds).
Symp.
Zool.
Soc.
London,
67:
277-289.
Fen on,
M.B.
and
Bell,
G.P.
(1981).
Recogni ion
o
species
o
insec i o ous
ba s
by
hei
echoloca ion
calls./.
Mamm.,
62:
233-243.
Fen on,
M.B.
and
Fulla d,
J.H.
(1979).
The
in luence
o
mo h
hea ing
on
ba
echoloca ion
s a egies./.
Comp.
Physiol.
A,
132:
77-86.
Fen on,
M.B.
and
Fulla d,
J.H.
(1981).
Mo h
hea ing
and
he
eeding
s a egies
o
ba s.
Am.
Scien is ,
69(3):
266-275.
Findley,
J.S.
(1993).
Ba s:
A
Communi y
Pe spec i e.
Camb idge
Uni e si y
P ess,
Camb idge.
Fluckige ,
P.F.
and
Beck,
A.
(1995).
Obse a ions
on
he
habi a
use
o
hun ing
by
Pleco us
aus-
iacus
(Fische ,
1829).
Myo is,
32—33:
121—122.
Fo es ,
T.G.,
Read,
M.P.,
Fa is,
H.E.
and
Hoy,
R.R.
(1997).
A
ympanal
hea ing
o gan
in
sca ab
bee les./.
Exp.
Biol.,
200:
601-606.
F ase,
B.A.,
Pizzu o,
T.M.
and
Ge z,
L.L.
(1990).
Su i o ship
o
bled
oles
measu ed
by
ecap-
u e./
Mamm.,
71:
104-105.
F eeman,
P.W.
(1981).
Co espondence
o
ood
habi s
and
mo phology
in
insec i o ous
ba s./.
Mamm.,
62(1):
166—170.
F os ,
D.R.
andTimm,
R.M.
(1992).
Phylogeny
o
pleco ine
ba s
(Chi op e a:
Vespe ilionidae):
summa y
o
he
e idence
and
p oposal
o
a
logis ically
consis en
axonomy.
Am.
Mus.
Na i a es,
3034:
1—16.
Fuh mann,
M.
and
Godmann,
O.
(1991).
Na u liche
qua ie e
de
wald lede emause
schu zen!
Konsequenzen aus
eine
Baumhohlenun e suchung
in
Rheingau.
AFZ-Allgemeine-Fo s
Zei sch i ,
49(19):
982-983.
Fuh mann,
M.
and
Sei z,
A.
(1992).
Noc u nal
ac i i y
o
he
b own
long-ea ed
ba
(Pleco us
au i us
L.1758):
da a
om
adio acking
in
he
Lennebu g
o es
nea
Mainz
(Ge many).
In:
Wildli e
Teleme y.
Remo e
Moni o ing
and
T acking
o
Animals
(P iede,
I.G.
and
Swi ,
S.M.,
eds).
Ellis
Ho wood,
Chiches e :
538-548.
Fulla d,
J.H.
(1987).
Senso y
ecology
and
neu oe hology
o
mo hs
and
ba s:
in e ac ions
in
a
global
pe spec i e.
In:
Recen
Ad ances
in
he
S udy
o
Ba s
(Fen on,
M.B.,
Racey,
P.A.
and
Rayne ,
J.M.V.,
eds).
Camb idge
Uni e si y
P ess,
Camb idge:
244-272.
Re e ences
161
Fulla d,
J.H.
(1992).
The
neu oe hology
o
sound
p oduc ion
in
ige
no hs
(Lepidop e a,
A c iidae).
1.
Rhy hmici y
and
cen al
con ol./.
Comp.
Physiol
A,
170.
575
5
.
Fulla d,
J.H.
and
Yack,
J.E.
(1993).
The
e olu iona y
biology
o
insec
hea ing.
I ee,
81/).
Funloslh,2K.
and
Uchida,
T.A.
(1982).
Annual
cycles
o
body
weigh
in
he
Namie's
os ed
ba
Vespe ilio
supe ans
supe ans.
J.
Zool.
London,
196:
417—430.
Gaisle ,
J.
(1966).
A
en a i e
ecological
classi ica ion
o
colonies
o
he
Eu opean
ba s.
Lynx,
Gaisk
J.
(1970)3Rema ks
on
he
he mop e e endum
o
palea c ic
ba s
in
hei
na u al
habi-
a s.
P oc.
2nd
In .
Ba
Res.
Con :
33-35-
Gaisle ,
J.
(1979).
Ecology
o
ba s.
In:
Ecology
o
Small Mammals
(S odda ,
D.M.,
ed.).
Chapman
&
Hall,
London:
281-342.
..
.
oq
nc
Gaisle ,
J.
(1983-4).
Ba s
o
no he n
Alge ia
and
hei
win e
ac i i y.
Myo is
21
22
89
99.
Gaisle ,
J.
and
Baue o a,
Z.
(1985-6). The
li e
o
ba s
in
a
ci y.
Myo is,
23-24:
211
215.
Gaisle ,
J.,
Chy il,
J.
and
Vlasin,
M.
(1990).
The
ba s
o
sou h
Mo a ian
lowlands
(Czechoslo akia)
o e
hi y
yea s.
P i odo edne
P ace
Us a u
Ceskoslo enske
Academie
ed
V
Ga i™?j24and
Hanak,
V.
(1964).
Ne opy
dlouhouchy
Pleco us
aus iacus,
Fische
1829
Bulha sku.
Zool.
Lis y,
1:
31-38.
.
^L„cim,0|,;a
Gaisle ,
J.
and
Hanak,
V.
(1969). Summa y
o
he
esul s
o
ba
banding
in
Czechoslo akia,
1948-1967.
Lynx,
10:
25-34.
,
Gaisle ,
J.,
Hanak,
V.
and
Ho acek,
I.
(1980-81).
Rema ks
on
he
cu en
s a us
o
ba
popula-
ions
in
Czechoslo akia.
Myo is,
18-19:
68-/5.
_
Gaisle ,
J.,
Zukal,
J.,
Nes adbo a,
J.,
Chy il,
J.
and
Obuch,
J.
(1996)
Species
di e si y,
and
el-
a i e
abundance
o
small
mammals
(Insec i o a,
Chi op e a,
Roden ia)
in
he
Pala a
bio-
sphe e
ese e
o
UNESCO.
Ac a
Soc.
Zool.
Bohem.,
60:
13-23.
.
Ga dne .
R.A.,
Molynenx,
D.H.
and
S ebbmgs.
R.E.
(1987).
S ud es
on
he
p e alence
o
haema ozoa
o
B i ish
ba s.
Mamm.
Re .,
17(2/3):
75-80.
Ge ell,
R.
(1980-81).
Ba
conse a ion
in
Sweden.
Myo is,
18-19:
H"15-
Ge ell,
R.
and
Lundbe g,
K.
(1985).
Social
o ganiza ion
in
he
ba
Pipis elluspipis ellus.
Beha .
Ecol.
Sociobiol.,
16:
177-184.
,
Glas,
G.H.
(1982).
Reco ds
o
hibe na ing
ba bas elle
and
g ey
long-ea ed
ba s
in
Ne he lands
ou side
he
sou he n
Limbu g
ca e
a ea.
Lu a,
25:
15-16.
Glue,
D.E.
(1970).
A ian
p eda o
pelle
analysis
and
he
mammalogis .
Mamm.
->3
Gould,
E.
(1955).
The
eeding
e iciency
o
insec i o ous
ba s.
J.
Mamm.,
36:
399-406.
Gould,
E.
(1971).
S udies
o
ma e nal-in an
communica ion
and
de elopmen
o
ocaliza ion
in
he
ba s
Myo is
and
Ep esicus.
Communica ions
Beha .
Biol.,
5:
263-313.
G eenaway,
F.
and
Hu son,
A.M.
(1990).
A
Field
Guide
o
B i ish
Ba s.
B uce
Coleman,
Uxb idge,
UK.
G i in,
D.R.
(1958).
Lis ening
in
he
Da k.
Yale
Uni e si y
P ess,
New
Ha en,
G1.
Gus a son,
A.W.
and
Damassa,
D.A.
(1987).
Binding
o
sex
s e oids
o
plasma
p o eins:
ela ion
o
and ogen
esis ance
and
asynch onous
ep oduc i e
pa e ns
in
hibe na ing
ba s.
In:
Recen
Ad ances
in
he
S udy
o
Ba s
(Fen on,
M.B.,
Racey,
P.A.
and
Rayne ,
J.M.V.,
eds).
Camb idge
Uni e si y
P ess,
Camb idge:
442-458.
Haensel,
J.
(1994).
Zum
Flede maus-Win e bes and
zahl eiche
in
de
S ad
Ba u h
o han-
dene' ,
eils
e allsge ah de e
E dkelle
-
Vo abei
u
ein
Schu zp og amm.
Nyc alus,
Be lin,
5(3/4):
249-273.
. .
.
Haensel,
J.
and
Na e,
M.
(1993).
Fla ismus
bei
einem
B aunen
Langoh
{Pleco us
au i us)
e he-
Index
Page
numbe s
in
i alics
e e
o
illus a ions.
accesso y
glands
70-/1
asynch onous
ec udescence
71
Accipi e
nisus,
p eda ion
by
117
ac i i y
58-59
e ec s
o
en i onmen al
ac o s
59
in
lac a ion
58-59
o
mo hs
58
ime
o
e u n
o
he
oos
58
unimodal
pa e ns
58
ae ial
insec
cap u e
39
A ican
alse
ampi e
ba
See
Ca diode ma
co
ageing, signs
o
108
agg ession
ho mones
esponsible
o
70
in e speci ic
di e ences
21,
104-105
in aspeci ic
21,
64,
101,
104
agili y
27
ai
mo emen s
in
oos s
15,
93
ai
empe a u e
e ec
on
ac i i y
59
e ec
on
eme gence
56-57
h eshold
o
insec
ligh
37,
128
Allen's
big-ea ed
ba
See
Idionyc e isphyllo is
al i ude
e ec
on
dis ibu ion
8-9
o
hibe nacula
123-124
ampulla y
glands
70
and ogen
esis ance
71
Anobium
punc a um
138
An ozous
pallidas
94
aposema ic
signals
34
A c iidae
in
die
43,
46
wa ning
clicks
34
a ousal
124-127
ene gy
expendi u e
129-130
equency
126-127
ac ile
and
non- ac ile
s imuli
129,
130
Asio
o us,
p eda ion
by
114
aspec
a io
27
asymp o ic
mass
81
baby
ba s
78-84
de elopmen
a
bi h
79-80
g ow h
a e
80-83
hand
ea ing
152
li e
size
78-79
pos -na al
de elopmen
80-84
size
a
bi h
78
he mo egula ion
84
anspo
in
ligh
79,
103
See
also
mo he -young
in e ac ion
baculum
16,
70
bank
ole
See
Cle h ionomys
gla eolus
Ba bas ella
9
Ba bas ella
ba bas ellus
ea
leng h
12
hibe na ion
empe a u e
123
ka yo ype
25
ba bed
wi e
133
ba
boxes
89,
97,
143-146
conc e e-sawdus
143,
145,
145
146
e ec i eness
o
schemes
143,
144,
146
use
by
long-ea ed
ba s
144-145
Ba
Conse a ion
T us
1,
153
ba
de ec o s
12,
30,
38,
109
ba
g oups
135,
147
ba
ings
51,
100,
108,
134
ing damage
137
Bechs ein's
ba
See
Myo is
bechs eimi
bee les
(Coleop e a)
in
die
41,
43—47
175

176
Index
black-headed
gull
See
La us
idibundus
booklice
(Psocop e a)
in
die
47
caddis
lies
(T ichop e a)
in
die
43—47
calcium,
e ec
on
li e
size
79
calo ime y
85-86
cap i e
ba s
152
Ca diode ma
co
37
ca ,
domes ic
See
Felis
ca us
ca e pilla s
in
die
37,
46
cauda
epididymis
72
cella s,
impo ance
as
hibe nacula
121-122,
147
cen ipedes
(Chilopoda)
in
die
37,
44,
45,
47
cha e s
(Sca abaeidae)
in
die
46,
47
chemical
ligh s
52
chi inase
49
ch omosomes
25
ci il
wiligh
54-55,
58
cladis ic
analysis
9
Cle h ionomys
gla eolus
119
Cloeo us
pe ci ali
36
cloud
co e
57
clus e ing
See
he mo egula ion
cobwebs
15,
93
colou
ace
18
pelage
18
a ia ion
along
hai s
18,
20
communica ion
calls
30,
83,
100
See
also
mo he -young
in e ac ion
condylobasal
leng h
17,
18
co pus
lu eum
74
Co yno hinus
mexicanus
9,
10
Co yno hinus
a inesquii
9,
10
age
o
sexual
ma u i y
71
Co yno hinus
ownsendii
9-10
audi o y
e iciency
23-24
clus e ing
94
ligh
dis ance
111
ges a ion
leng h
77
hibe na ion
empe a u e
123
pos -na al
de elopmen
80,
82
age
o
sexual
ma u i y
71,
75
Co yno hinus
ownsendii
ingens
10
ac i i y
pa e ns
59
ime
o
eme gence
55
Co yno hinus
ownsendii
i ginianus
9—10
ac i i y
pa e ns
59
e ec s
o
ansmi e s 54
adio
acking
52,
54
Cowpe 's
glands
69,
70
c ane lies
(Tipulidae)
in
die
43,
46
c eches
80
c op
sp aying
133
dac ylopa agium
26
dac ylopa agium
mino
26
dayligh
lying
e ec
o
en i onmen al
ac o s
116,
128
p e alence
116-117
easons
o
117-118
dea h
wa ch
bee le
See
Xes obium
u o illosum
deciduous
ees
impo ance
o
o aging
61,63
p oximi y
o
oos s
64,
92
dehyd a ion
50,
127
delayed
o ula ion
68
den i ion
See
ee h
die
P.
au i us
43—46
P.
aus iacus
46-47
diges i e
e iciency
49
dis ibu ion
P.
au i us
3-9,
4,
6
P.
aus iacus
3-9,
5,
7
dis u bance
in
hibe nacula
129-131,
134,
136
in
summe
oos s
93, 95,
134,
136
doubly
labelled
wa e
(DLW)
echnique
85-86,
87
e ec
on
ba s
137
d oppings
colou
15
dis ibu ion
in
oos s
15,
135
size
15
olume
in
oos s
93,
135
d owning,
dange
o
135
ea s
22-24
d ag
e ec
24,
28
e ec ion
1
olding
1, 12,
14,
22-23,
131
unc ions
23-24
leng h
2,
12
medial
lobules
22
am's
ho n
12,
14,
22—23
sensi i i y
24,
38
s uc u e
22
he mo egula o y
unc ion
23
ea wigs
(De map e a)
in
die
43—46,
47
eas e n
big-ea ed
ba
See
Co yno hinus
a inesquii
echoloca ion
30-32
de ec ion
dis ance
13
ene ge ic
cos
32
equency
13,
30-31
in ensi y
o
calls
12,
36,
60,
64,
109
loud,
long-sweep
30,
31,
100
p oduc ion
h ough
nose
25
pulse
a e
30-31
ec opa asi es
leas
112
in es a ion
le el
112
mi es
112
nyc e ibiid
lies
112
eme gence
beha iou
56
clumped
56
e ec
o
en i onmen al
ac o s
56—57
ime
o
54-55,
37-58,114
emig a ion
100-101,
106
endopa asi es
112
endo he my
87
ene gy
cos
o
ligh
86
demand
in
lac a ion
85
demand
in
p egnancy
85
equi emen
o
milk
p oduc ion
85
epididymis
70
Ep esicus iuscus
84
acial
glands
103
Ep esicus
nilssonii,
die
48
Ep esicus
se o inus
eyesigh
24
acial
glands
103
ime
o
eme gence
54
Eude ma
macula um
9,
10-11
o aging
beha iou
11,
35—36
Eu opean
Ba Resea ch
O ganiza ion
1
exclusion
o
colonies
133,
135,
147
eyesigh
24
acial
glands
103,
104
aecal
analysis
40-41,
43-47
Falco
innunculus,
p eda ion
by
114,
117
ecundi y
77-78
eeding
pe ches
1,
15,
40, 43,
90
eeding
a e
66-67
eeding
si es
62-64
densi y
o
ba s
63-64
equency
o
use
63
size
63
Felis
ca us,
p eda ion
by
115-116,
117
emale
de ence
polygyny
104,
105
e al
g ow h
a e
77
issu e
oos ing
90,
91
lies
(Dip e a)
in
die
44,
4}-47
ligh
cha ac e is ics
in
Pleco us
13
dis ances
110-111
li
coe icien
29
minimum
powe
speed
28
speed
27-28,
60
s aigh
ligh
28
unnel
29
u ning
ligh
28-29
wi hin
oos s
See
oos s
See
also
ho e ing
lyways
59-60,
62,
133,
146
og,
e ec
on
o aging
59
o aging
dis ance
64-66,
101
du a ion
67,
86
eeding
si es
62-64
habi a
60-62
,,
impo ance
o
deciduous
woodland
61,
14b
inside
oos s
46,
60-61,
128
o ea m
b oken
151
leng h
19
a e
o
g ow h
in
ju eniles
82
ges a ion
76-77
delayed
76,
77
e ec
o
en i onmen al
ac o s
/b,
/ /
leng h
77
gleaning
36-39,
56
ad an ages
37,
58,
128
echoloca ion
du ing
31-32,
38
g ound
gleaning
36,
45
ho e ing
du ing
29-30,
37
p opo ion
o
die
gleaned
45-47,
48,
62
gonado ophins
76
G ahamella
sp.
112
habi a
des uc ion
133
pa chiness
60
handling
ime
86
ha es men
(Opiliones)
in
die
3/,
hea ing
See
ea s
he e o he my
119
hibe nacula
120-124
ai
low
122
al i ude
123-124
178
Index
hibe nacula
con .
a i icial
147-148
humidi y
123
sinan h opic
si es
121-122,
132
si e
changing
122
empe a u e
122-123
hibe na ion
cha ac e is ics
119-120
posi ions
124,
125,
126
si es
121-124
iming
120
weigh
loss
121
hi undine
bi ds,
in e ac ion
wi h
ba s
117
his oplasmosis
136
homing
111
ho e ing
29-30,
38-39
dis ance
39
i-calls
80,
83-84,
102,
103
pos -na al
changes
83-84
See
also
mo he -young
in e ac ion
Idionyc e isphyllo is
9,
10
inju ies
116,
134,
151
insec
emains
benea h
eeding
pe ches
15,
40,
43
in
aecal
pelle s
40-41
Ischnopsyllus
hexa inus
112
jaw
mo phology
17-18,
48
ka yo ypes
See
ch omosomes
kes el
See
Falco
innunculus
lacewings
(Neu op e a)
in
die
43-45,
47
esponses
o
ba
calls
35
ympanic
o gans
34-35
lac a ion
du a ion
84
e ec
on
ac i i y
pa e ns
58-59
e ec
on
o aging
beha iou
64-65
See
also
ene gy
La us
idibundus,
p eda ion
by
117-118
Lasiu us
cine eus
35
La ia
ons
104
legal
p o ec ion
134-135
leks
104
Lep o ombidium
ussicum
112
li e
expec ancy
106,
107
li e
span
a e age
106,
107
maximum
108
ligh
sampling
beha iou
55,56
ligh
aps
41
lindane
138,
139,
140
li e
size
78-79
cons ain s
on
79
longe i y
108
lump-nosed
ba s
See
Co yno hinus
sp
Malaise
ap
41,
42
male
colonies
99,
100-101
mandible
leng h
17
mandibula
oo h
ow
18
manoeu abili y
27,
29
ma ing
68,
69,
73-74
sp ing
73,
104
swa ming
104—105
sys ems
74,
103-105
win e
73,
104
maxilla y
oo h
ow
17-18
may lies
(Epheme op e a)
in
die
41
Melolon ha
in
die
47
Mexican
big-ea ed
ba
See
Co yno hinus
mexicanus
midges
(Nema oce a)
in
die
45
mig a ion
119
Miniop e us
sch eibe sii
101
mobbing
by
bi ds
117
monogamy
104
mo ali y
ju enile
84
a e
106,
108,
115
mo he -young
in e ac ion
abandonmen
108
acous ic
cues
102-103
ol ac o y
cues
102-103
ecogni ion
102-103
e ie al
103
eunion
beha iou
102
selec i e
nu sing
101-102
mo hs
(Lepidop e a)
abundance
45,
61
in
die
43—47
lu e ing
38,
46
emains
benea h
eeding
pe ches
43
esponses
o
ba
calls
34
ympanic
o gans
33—34
mouse-ea ed
ba
See
Myo is
myo is
Myo is
bechs einii
die
49
ea
leng h
12
Index
179
Myo is
bly hii
40
Myo is
dasycneme
138
Myo is
dauben onii
die
48
diges i e
e iciency
49
dis u bance
in
oos s
93
ligh
speed
28
use
o
lyways
60
wa e
balance
50
Myo is
e o is
37
Myo is
luci ugus
a ousal
equency
127
audi o y
e iciency
23-24
echoloca ion
36
ene ge ics
o
ep oduc ion
37
Myo is
myo is
36,
40
delayed
ges a ion
76
de elopmen
in
ju eniles
84
p eda ion
by
owls
114
su i al
a e
108
Myo is
na e e i
die
36,45,
48-49
dis u bance
in
oos s
93
echoloca ion
calls
13,
36
ligh
speed
28
gleaning
36,
45,
49
use
o
lyways
60
Myo is
sep en ionalis
36
Myo is
sodalis
111
Na e e 's
ba
See
Myo is
na e e i
Neomyobia
pleco ia
112
nigh
oos s
See
eeding
pe ches
Noc uidae
in
die
43, 46,
esponses
o
ba
calls
34,
ympanic
o gans
33,
34
nos ils
p oduc ion
o
calls
25
shape
25
nu se y
colonies
cohesion
99,
100
composi ion
99-100
ac o s
a ec ing
colony
size
98,
100
p esence
o
males
99,
100-101
size
97-98
Nyc alus
noc ula
age
o
sexual
ma u i y
75
ligh
speed
28
ma ing
104
mig a ion
111,
119
spe m
s o age
73,
75
ime
o
eme gence
54
Nyc e ibia
schmidlii
112
Nyc e idopsylla
longiceps
112
oppoXis i
o aging
39-40,
45, 46,
61,
64
o ganochlo ines
133,
138—140
o ien a ion
calls
83,
84
o igins
P.
au i us
3
P.
aus iacus
3
O ni honyssus
pipis elli
112
ou bu s
beha iou
56
o a ies
74
o e hea ing
117
o ula ion
76
owls
ba n
See
Ty o
alba
long-ea ed
See
Asio
o us
pelle s
114
popula ion
size
115
awny
See
S ix
aluco
Oza k
big-ea ed
ba
Co yno hinus
ownsendu
ingens
pa u i ion
da e
77
posi ion
78
passi e
lis ening
38-39,
46
penis
shape
20
pen achlo ophenol
(PCP)
138,
139,
140
pe cen age
equency
( aecal
analysis)
41
pe cen age
occu ence
( aecal
analysis)
4
pe ch
hun ing
37
pe me h in
138—139,
140
pho og aphy
inside
oos s
134
phylogene ic
echniques
9
pipis elle
See
Pipis ellus
pipis ellus
Pipis ellus
pipis ellus
ac i i y
pa e ns
37,
58
age
o
sexual
ma u i y
75
delayed
ges a ion
76-77
eme gence
beha iou
56
ligh
29
ma ing
104
mig a ion
119
milk
p oduc ion
87
plasma
p oges e one
le el
74
oos
si es
89,
93
spe m
s o age
75
ime
o
eme gence
54
180
Index
Pipis ellus
pipis ellus
con .
win e
lying
127
placen a,
ea ing
o
78
plagiopa agium
26,11
pleco ine
g oup
1,
9-11
Pleco us
au i us
wa di
2
Pleco us
aus iacus
kolomba o ici
8
Pleco us
b e imanus
2
Pleco us
mexicanus
See
Co yno hinus
mexicanus
Pleco us
phyllo is
See
Idionyc e is
phyllo is
Pleco us
a inesquii
See
Co yno hinus
a inesquii
Pleco us
ai anus
2
Pleco us
ene i ae
2
Pleco us
ownsendii
See
Co yno hinus
ownsendii
poisoning
See
emedial
imbe
ea men
popula ion
densi y
109
size
108-110
pos e
campaigns
109
p eda ion
ae ial
114-115,
117-118
e ec
on
mo ali y
a e
115
in luence
on
ba
beha iou
113
mammalian
115-116
isk
55,
60,
114,
118
p eda o
a oidance
beha iou
56,
117-118,
131
p e-hibe nal
weigh
gain
120—121
p ey
a ailabili y
40,
58
p ey-gene a ed
sounds
38-39,
46
p oges e one
74
p opa agium
26
p os a e
gland
70
abies
136
adian
hea ,
e ec
on
body
empe a u e
117
adio
eleme y
52-54,
127,
134
adio
ansmi e s
a achmen
me hods
52-53
e ec s
on
ba
beha iou
54,
136-137
Ra inesque's
big-ea ed
ba
See
Co yno hinus
a inesquii
ain
e ec
on
ac i i y
59
e ec
on
eme gence
56
amus
mandibulae
17
ecip ocal
al uism
94
ecogni ion
be ween
colony
membe s
100
o
mo he s
and
in an s
See
mo he -young
in e ac ion
e lec i e
ape
50,
60
ela i e
body
condi ion
71
emedial
imbe
ea men
chemicals
used
138-139
legal
posi ion
134,
140
oxici y
o
ba s
139-140
ep oduc i e
o gans
emale
74
male
69-70
ep oduc i e
a e
69
esou ce
de ence
polygyny
104
esou ce
pa i ioning
48-49
Rhinolophidae
25,
75
Rhinolophus
24,
29
Rhinolophus
e umequinum
de elopmen
in
ju eniles
84
echoloca ion
36,
40
eme gence
beha iou
57
ligh
29
selec i e
o aging
40
Rhinolophus
hipposide os
delayed
ges a ion
76
echoloca ion
36
Rhizo agus
aes i us
46
oos
coun s
98,
109
oos ing
posi ions
90,
91
oos s
age
92-93
des uc ion
133
ideli y
o
93,
97,
100,
101
lying
inside
90,
92-93,
135-136
legal
p o ec ion
134—135
ma king
o
104—105
ma ing
104—105
nu se y
68,
90
signs
o
15
empo a y
90, 95,
109,
112
ansi o y
97,
104
oos
selec ion
90-92
habi a
92
physical
ea u es
92-93
p oximi y
o
woodland
92,
93
wa m h
90, 92,
93
oos
si es
chu ch
lo s
90
a m buildings
90, 95,
147
houses
89,
90,
92-93,
132, 133,
ee
holes
89,
90, 95,
97,
121,
146
oos
empe a u e
90, 93,
94,
95
Roos
Visi o 's
licence
134
Sca hop
haga
s e co
a ia
37,
45
scen
in
mo he -young
ecogni ion
102-103
Index
181
in
ecogni ion
o
colony
membe s
100
selec i e
o aging
40,
47
selec i e
nu sing
See
mo he -young
in e ac ion
se o ine
See
Ep esicus
se o inus
sex
a io
110
sex
s e oid
binding
ho mone
(SBP)
71
sexual
dimo phism
18-19
sexual
ma u i y
age
o
71,
74-75
ac o s
a ec ing
71-72
me hods
o
assessing
72,
75,
110
si es
o
special
scien i ic
in e es
(SSSIs)
134-135
skull
16-18,
17
measu emen s
o
sepa a ing
species
17-18
mo phology
48
social
acili a ion
64
space
oos ing
90,
91
spa owhawk
See
Accipi e
nisus
spe m
s o age
by
males
73
s o age
in
u e us
68,
73,
75-76
spe ma ogenesis
68,
69,
72-73
e ec
o
en i onmen al
ac o s
72
e ec
o
o po
72-73
ho monal
con ol
70-71
spide s
(A aneae)
in
die
37,
44,
45,
46,
47
Spinc u nix
pleco ina
112
spo ed
ba
See
Eude ma
macula um
s a u o y
na u e
conse a ion
bodies
134
s anded
ba s,
ca e
o
151
s ee
lamps
exploi a ion
by
ba s
47,
140-142
ypes
o
lamp
140
S ixaluco,
p eda ion
by
114
suckling
au oc ine
eedback
59
equency
58-59
suc ion
aps
41,
45
supe s i ions
135
su i al
a e
106-107
swa ming
104-105
sweep
ne
43,
61
swi ching
ood
sou ces
119
Tada ida
b asiliensis
101
Taiwan
long-ea ed
ba
See
Pleco us
ai anus
ee h
20-21,21
canine
leng h
20,
48
den al
o mula
20
pa ame e s
o
di e en ia ion
o
species
20
p emola s
20,
21
Teneb io
sp
49,
152
Tene i e
long-ea ed
ba
See
Pleco us
ene i ae
e i o iali y
101
es es
70-71
asynch onous
ec udescence
71
he mo egula ion
clus e ing
94-95,
101
du ing
p egnancy
and
lac a ion
86-87,
93-94
du ing
spe ma ogenesis
72-73,
93-94
a high
empe a u e
95,
96
humb
19
leng h
19
leng h/ o ea m
a io
19
o po
daily
67,
119-120
e ec
on
milk
p oduc ion
87-88
ex ended
119,
128
in
ep oduc ion
86—88
in empo a y
oos s
95-97
Townsend's
big-ea ed
ba
See
Co yno hinus
ownsendii
agus
20,
22
shape
20
wid h
19-20
ue
bugs
(Hemip e a)
in
die
45-47
ympanic
o gans
33,
35
unica
aginalis
72
winning
a e
78-79
ympanic
bullae
17,
18
ympanic
o gans
e olu ion
in
insec
g oups
33,
34-35
ange
o
hea ing
35
s uc u e
33-34
Ty o
alba,
p eda ion
by
114
u baniza ion
92,
143
u e h a
70
u opa agium
26,
39
u e us
75-76
u iliza ion-a ailabili y
es
61,63
asa
e e en ia
70
as
de e ens
70,
73
Vi ginia
big-ea ed
ba
See
Co yno hinus
ownsendii
i ginianus
ision
24
in
homing
111
ocaliza ions,
pos -na al
83-84
182
Index
wasps
(Hymenop e a)
in
die
45-47
wa e
balance
49-50
wa e
loss
e apo a i e
50
aecal
50
in
u ine
50
wa e
anks
135
weaning
84
weigh
a
bi h
78
inc ease
du ing
p egnancy
86,
87
loss
due
o
dis u bance
in
hibe nacula
129-130
loss
du ing
hibe na ion
121
loss
du ing
p e- ligh
pe iod
84
a e
o
pos -na al
inc ease
81-82
wes e n
big-ea ed
ba
See
Co yno hinus
ownsendii
whispe ing
ba s
12,
30
Wildli e
and
Coun yside
ac
1981
134,
140
wing
a ea
27
cambe
28
ac u es
151
loading
27,
29,
30,
54
mo phology
26
pos -na al
de elopmen
82-83
shape
26-27,
30
win e
ligh s
e ec
o
ai
empe a u e
128-129
ood
consump ion
128
equency
128
pu pose
127-128
iming
128
woodland
coni e ous
60-62,
133,
143
deciduous
60-62,
64,
92,
132,
133
edges
61,
133
e ec
on
colony
size
98
woodwo m
See
Anobium
punc a um
Xes obium
u o illosum
138