Ecology of European bats in relation to their conservation

Ba
Biology
and
Conse a ion
Edi ed
by
Thomas
H.
Kunz
and
Paul
A.
Racey
SMITHSONIAN
INSTITUTION
PRESS
Washing on
and
London
248
E.
D.
PIERSON
AND
P.
A.
RACEY
ba
con ol
in
Middle
and
Sou h
Ame ica,
and
killing
o
ood
o
because
o
misconcep ions
ega ding
disease
isks
in
A ica,
in
Asia,
and
on
Paci ic
islands.
The
endency
o
ca e-dwelling
ba s
o
o m
la ge
agg ega ions
and
be
highly
isible,
howe e ,
ende s
hem
pa icula ly
ulne able
o
human-induced
dis u bance
o
mo ali y.
Conse a ion
e o s
a e
equen ly
hampe ed
by
in-
su icien
knowledge
o
he
dis ibu ion
and
na u al
his o y
o
mos
species.
Dis ibu ional
maps
a e
gene ally
a
be e
indica ion
o whe e
s udies
ha e
been
conduc ed
han
whe e
he
ba s
occu :
New
dis ibu ion
eco ds
a e
s ill
eme ging,
e en
in
egions
ha
ha e
been
in ensi ely
s udied.
Addi ion-
ally,
he
speci ic
ecological
in o ma ion
sough
by
land
man-
age s
is
gene ally
una ailable—in o ma ion
such
as
he
numbe
o
dead
oos
ees
pe
hec a e
ha
should
be
e-
ained
in
imbe
ope a ions,
he
mic oclima ic
condi ions
ha
de e mine
oos
selec ion,
he
lexibili y
o
a
species
in
i s
oos ing
o
o aging
equi emen s,
and
he
ex en
o
which
habi a
al e a ion
a ec s
longe i y
o
ep oduc i e
success.
Un esol ed
axonomic
issues
can
also
inhibi
he
o mu-
la ion
o
species-speci ic
conse a ion
s a egies.
F om
bo h
he
biological
and
he
poli ical
pe spec i e
i
is
necessa y
o
de ine
he
axonomic
uni
o
which
a
policy
is
ecom-
mended.
A
axon
may
be
assigned
di e en
conse a ion
p io i ies
depending
on
whe he
i
is
iewed
as
a
subspecies
o
a
dis inc
species,
and
i
can
be
o e looked
al oge he
i
i s
co ec
axonomic
s a us
is
no
ecognized.
New
species
a e
s ill
being
disco e ed,
iden i ied,
o
ede ined
e en
in
a eas
such
as
Eu ope
and
No h
Ame ica,
which
ha e
been
sub-
jec
his o ically
o
ela i ely
close
axonomic
sc u iny.
Pub^
misconcep ions
ega ding
ba s
ha e
hwa ed
con-
se a ion
e o s
e e ywhe e,
bu
he
sou ce
o
misunde -
s anding
can
di e
egionally.
In
No h
Ame ica
i
may
be
ea
o
abies,
in
Middle
and
Sou h
Ame ica
ea
o
ampi e
ba s,
o
in
Aus alia
conce n
o e
p eda ion
in
ui
o cha ds.
The
esul ,
howe e ,
is
he
same:
ba s
a e
o en
ea ed
as
e min,
subjec ed
o
e adica ion
campaigns,
and
o e looked
o
unde alued
by
land
manage s.
Mos
o
he
ollowing
chap e s
ecognize
wo
basic
ap-
p oaches
o
conse a ion:
p o ec ion
o
indi idual
species
and
p ese a ion
o
habi a .
Iden i ied
isk
ac o s
o
in-
di idual
species
include
small
popula ion
size
o
a i y,
e idence
o
popula ion
declines,
limi ed
dis ibu ion,
e-
s ic ed
habi a
equi emen s,
and
pa icula
ulne abili y
o
human-induced
impac s.
The
highes
p io i y
is
usually
g an ed
o
endemic
and
mono ypic
species.
A
species-
ocused
app oach
has
been
and
likely
will
be
c i ical
o
he
pe sis ence
o
ce ain
axa.
Fo
example,
in
a eas
whe e
ba s
a e
in ensi ely
hun ed
o
subjec ed
o
sys ema ic
e adi-
ca ion,
habi a
p o ec ion
does
no
su ice.
An
addi ional
app oach
is
o
p o ec
habi a s
o
habi a
ea u es
c i ical
o
he
su i al
o
ba
communi ies.
Al-
hough
habi a -o ien ed conse a ion
e o s
gene ally
o-
cus
on
a eas
o
high
biodi e si y,
his
is
no
su icien
o
ba s.
Because
o
he
e y
p onounced
la i udinal
g adien
in
species
di e si y,
i
species
ichness
we e
he
only
c i e-
ion
many
habi a s
impo an
o
ba s
would
be
o e looked.
Because
ba
dis ibu ion
is
equen ly
mo e
closely
linked
o
s uc u al
a he
han
bo anical
cha ac e is ics
o
he
land-
scape,
conse a ion
s a egies
based
solely
on
bo anical
communi ies
may
no
accoun
o
he
habi a
ea u es
mos
needed
by
many
ba
species
(e.g.,
ca es
o
ock
o ma ions).
In
opical
egions,
low-ele a ion
and
coas al
o es s
ha e
been
mos
ulne able
o
con e sion o
human
de elop-
men .
The
loss
o
d y
o es s,
which
do
no
ecei e
he
same
conse a ion
a en ion
as
ain o es s,
is also
o
se ious
conce n.
The
au ho s
ha e
gene ally
ecognized
ha
conse a ion
s a egies
o
any
species
o
habi a
mus
include
p o ec ion
o
bo h
oos ing
and
o aging
a eas.
Acknowledgmen
o
oos ing
equi emen s
gene ally
leads
o
he
ecogni ion
ha
ca e ns
(ca es,
old
mines,
and
o he
an h opogenic
s uc u es)
and
ma u e
na i e
o es s
a e
c i ical
o
he
p o-
ec ion
o
bo h
numbe s
o
species
and
o e all
di e si y.
Fo aging
needs
a e,
in
gene al,
less
well
unde s ood
han
oos ing
equi emen s,
bu
unpollu ed
wa e
sou ces,
in ac
ipa ian
zones,
complex o es
s uc u e,
and
subs an ial
ac s
o
na i e
habi a
a e
men ioned
by
a
numbe
o
au ho s
as
impo an
o
he
main enance
o
sui able
o ag-
ing
a eas.
While
he
mos
e ec i e
conse a ion
ac ions
a e
o en
based
on
g ass- oo s
e o s
and
equi e
he
dedica ion
o
many
people
a
a
local
le el,
na ional
and
egional
policies
p o ec ing
ba
popula ions
a e
also
necessa y.
The
Aus a-
lian
Ba
Ac ion
Plan
(Richa ds
and
Hall),
he
es ablishmen
o
ede al
conse a ion
uni s
in
B azil
(Ma inho-Filho
and
Sazima),
he Eu opean
Ba s
Ag eemen
(Racey),
he
Uni ed
S a es
Endange ed
Species
Ac
(Pie son),
o
CITES
(Con-
en ion
on
In e na ional
T ade in
Endange ed
Species
o
Wild
Fauna
and
Flo a)
egula ions
(Rainey)
se e
as
cases
in
poin .
Pa
Fou
ep esen s
a
signi ican
benchma k
as
he
i s
symposium
o
ocus
on
he
conse a ion
biology
o
ba s.
The
inclusion
o
his
opic
in
a
olume
o
in e na ional
scope
signals
he
in en ion
o
ba
biologis s
o
p o ide
a
scien i ic
basis
o
he
conse a ion
o
hei
s udy
animals,
so
as
o
in o m
policy
make s
and
manage s,
in
he
hope
ha
ba s
will
con inue
o
be
majo
con ibu o s
o
mam-
malian
biodi e si y
wo ldwide.
ELIZABETH
D.
PIERSON
AND
PAUL
A.
RACEY
17
Ecology
o
Eu opean
Ba s
in
Rela ion
o
Thei
Conse a ion
PAUL
A.
RACEY
Be o e
ba
de ec o s
enabled
ield
iden i ica ion
o
o aging
ba s
(Ahlen
1981;
Baagoe
1986,
in
manusc ip ),
he
majo i y
o
s udies
o
Eu opean
species
in ol ed
coun ing
indi id-
uals
and
desc ibing
hei
ac i i y
and
oos ing
habi s
in
ma e ni y
colonies
and
hibe nacula.
G ea
dispa i ies
we e
o en
seen
be ween
summe
and
win e
coun s,
pa icula ly
o
espe ilionids,
wi h
win e
popula ions
accoun ing
o
a
small
p opo ion
o
hose
coun ed
in
summe
and
ice
e sa,
depending
on
species.
Al hough
he
mos
eliable
es ima es
o
popula ion
size
ha e
gene ally
been
de i ed
om
s udies
o
ma e ni y
colonies
(Gaisle
1975,
1978;
Speakman
e
al.
1991a;
En wis le
1994),
he e
emains
con-
side able
di icul y
in
es ablishing
eliable
popula ion
ends
in
Eu opean
ba
species.
Conse a ion
legisla ion
in
many
Eu opean
coun ies
has
p o ec ed
ba s
bu
no
hei
oos s
and
habi a s
(S eb-
bings
1988).
Mo e
ecen ly,
howe e ,
he
Eu opean
Ba s
Ag eemen
1992
(pa
o
he
1979
Bonn
Con en ion
on
he
Conse a ion
o
Mig a o y
Species
o
Wild Animals)
and
he
Eu opean
Union
(EU)
Habi a s
and
Species
Di ec i e
1992
ha e
di ec ed
a en ion
owa d
he
p ese a ion
o
o aging
habi a s.
Al hough
he
minia u iza ion
o
a-
dio ansmi e s
and
he
applica ion
o
adio acking
in
e-
cen
yea s
has
aided
he
s udy
o
ba
habi a
equi emen s,
gene aliza ions
applicable
on
a
na ionwide
scale
ha e
no
hi he o
been
possible
in
any
Eu opean
coun y.
The
U.K.
Na ional
Ba
Habi a
Su ey
The
i s
na ional
ba
habi a
su ey
was
ini ia ed
in
he
Uni ed
Kingdom
in
1990,
and
he
esul s
ha e
been
published
by
Walsh
e
al.
(1995)
and
Walsh
and
Ha is
(1996a, 1996b).
The
su ey
adop ed
a
andom
s a i ied
sampling
sys em
(Magu an
1988)
based
on
a
land
classi ica ion
ha
assigns
e e y
1-km
squa e
in
B i ain
o
1
o
32
land
classes.
Squa es
in
each
land
class
ha e
a
simila
clima e,
physiogeog aphy,
and
pa e n
o
land
use
(Bunce
e
al.
1981a,
1981b,
1983).
Wi hin
each
land
class
a
sample
o
1
-km
squa es
was
selec ed
a
andom,
o
a oid
obse e
bias
in
he
selec ion
o
si es
and
o
ensu e
a
s anda d
sampling
e o
in
di e en
landscape
ypes.
Fieldwo k
was
ca ied
ou
du ing
h ee
consecu i e
sum-
me s
om
1990
o
1992
and
in ol ed
bo h
p o essional
and
ama eu
ba
biologis s,
he
la e
d awn
mainly
om
B i -
ain's
90
ba
g oups.
Each
olun ee
was
alloca ed
one
o
mo e
1-km
squa es
and
walked
a
ansec
in
each
squa e
249
250
P.
A.
RACEY
ou
imes
du ing
p ede e mined
pe iods
in
summe ,
a oid-
ing
nigh s
when
wea he
condi ions
we e
un a o able
o
ba s.
Volun ee s
ca ied
unable
ba
de ec o s
(mainly
QMC
Mini-2)
se
a
45
kHz
o
maximize
he
ange
o
species
encoun e ed,
and
no ed he
o al
numbe
o
ba
passes
and
eeding
buzzes
in
each
squa e
and
wi hin
each
habi a
ype.
The
mo e
expe ienced su eyo s
we e
able
o
iden i y
some
ba s
o
species
o
species
g oups.
Analysis
o
he
da a
e ealed
ela ionships
be ween
ba
ac i i y
and
habi a
a iables
wi hin
and
ac oss
he
32
land
classes,
which
o
he
pu poses
o
he
analysis
we e
com-
bined
in o
se en
majo
g oups:
h ee
a able,
wo
pas o al,
ma ginal
upland,
and
upland.
A oidance
o
selec ion
o
habi a
ypes
was
examined
by
cons uc ing
Bon e oni
con idence
in e als
a ound
he
obse ed
use
o
each
habi-
a
ype
(Neu
e
al.
1974),
and
eg ession
analyses
we e
used
o
e alua e
habi a s
o
c i ical
impo ance
in
de e mining
high
ba
ac i i y
in
each
land-class
g oup.
O
he
1,030
su eyed
1-km
squa es,
910
p o ided
da a
sui able
o
analysis,
in ol ing
2,700
hou s
o
sea ch
e o
wi h
30,000
ba
passes
eco ded
in
he
9,000
km
walked.
Obse e s
iden i ied
24%
o
ba
passes
o
a
pa icula
spe-
cies
o
species
g oup,
and
71%
o
hese
we e
Pipis ellus
pipis ellus,
17%
Myo is
spp.,
8%
Nyc alus
noc ula,
3%
Pleco us
spp.,
and
2%
Ep esicus
se o inus.
Because
a
simila
p opo -
ion
o
he
uniden i ied
ba
passes
we e
p obably
Pipis ellus
pipis ellus,
habi a
p e e ences
(and
hei
implica ions
o
conse a ion)
apply
o
he
Vespe ilionidae
as
a
whole
and
o
P.
pipis ellus
in
pa icula .
Land
class
was
demons a ed
o
be
a
highly
signi ican
ac o
in luencing
he
incidence
o
ba
passes,
wi h
he
g ea es
ba
ac i i y
occu ing
in
pas o al
land
classes.
Ac oss
all
land-class
g oups,
ba s
ended
o
o age
selec-
i ely
in
edge
and
linea
habi a s
and
a oided
mo e
open
and
in ensi ely
managed
habi a
ypes.
They
showed
a
a
s onge
p e e ence
o
all
bodies
o
wa e
and
woodland
edges
han
o
any
o he
habi a
ype,
emphasizing
he
impo ance
o
hese
habi a s
as
key
o aging
si es.
In
he
woodland
ca ego y,
edges
we e
mo e
s ongly
selec ed
han
openings,
and
semina u al
b oad-lea ed
woodland
was
mo e
s ongly
selec ed
han
ei he
mixed
o
coni e ous
woodland.
U ban
a eas,
which
included
illages,
small
owns,
and
he
subu bs
o
some
la ge
ci ies,
we e
selec ed
in
h ee
land-class
g oups.
Linea
ege a ion
co ido s,
pa -
icula ly
ee-lined
hedge ows
and
co e ed
di ches,
we e
also
selec ed
by
ba s,
emphasizing
he
impo ance
o
land-
scape
connec i i y.
Sandy,
shingle,
o
ocky
beaches
and
es ua ine
coas al
ma sh
we e
also
signi ican ly
selec ed
as
o aging
si es.
Habi a s
s ongly
and
consis en ly
a oided
in
all
land-class
g oups
we e
hose
ha
we e
mo e
exposed
and
mo e
in ensi ely
managed,
including
moo land,
im-
p o ed
g assland,
upland,
and
a able
land.
The
only
g ass-
land
ype
no
consis en ly
a oided
was
lowland,
unim-
p o ed
g assland.
Ba s
o aged
p e e en ially
in
habi a s
ha
we e
com-
pa a i ely
a e
wi hin
each
land-class
g oup.
Fo
example,
he
pe cen age
a ailabili y
in
each
land-class
g oup
o
he
p e e ed
habi a s
o
woodland
edge,
ee
lines,
hedge ows,
and
bodies
o
wa e
anged
om
14%
o
31%,
wi h
a
mean
o
25%.
In
con as ,
he
a ailabili y
o
habi a s
ha
we e
consis endy
a oided
(s one
walls,
moo land,
a able,
and
mos
g assland
ca ego ies)
anged
om
40%
o
54%,
wi h
a
mean
o
47%.
Op imal
habi a s
end
o
be
a
he
pe ime-
e s
o
o he
habi a
ypes
o
linea
s ips,
and
hus
in
compa ison
wi h
con iguous
blocks
o
pas u e
o
a able
land,
o
ins ance,
hei
a ea
is
p opo ionally
smalle .
Bod-
ies
o
wa e
gene ally
ep esen
less
han
1%
o
he
a ail-
able
habi a ,
and
b oad-lea ed
woodland
edge
anges
om
3%
o
4%.
Because
he
selec ion
pa e ns
we e
consis en
be ween
indi idual
land
classes,
he
esul s
o
he
analysis
by
Walsh
and
Ha is
(1996a,
1996b)
can
be
summa ized
by
habi a
ype
a he
han
by
land
class
(Table
17.1).
The
p ima y
aim
o
he
U.K.
Na ional
Ba
Habi a
Su -
ey
was
o
p o ide
a
means
o
assessing
he
signi icance
o
habi a
change
o
ba
popula ions.
By
expanding
he
scale
o
p e ious
s udies
o
habi a
p e e ences
( e iewed
in
Walsh
e
al.
1995)
and
dis ibu ion
su eys
(Ahlen
1980-
1981;
Baagoe
1986;
Rydell
1986;
Jiides
1989)
o
a
na ional
le el
and
using
a
land
classi ica ion
sys em
widely
accep ed
in
he
Uni ed
Kingdom,
Walsh
e
al.
(1995)
and
Walsh
and
Ha is
(1996a,
1996b)
ha e
de eloped
a
po en ial
me hod
o
de ec ing
change,
es ablishing
i s
di ec ion
and
measu ing
i s
magni ude
using
a
p o ocol
o
simul aneously
moni o -
ing
ba
ac i i y
and
habi a
use.
Analysis
o
habi a
ac o s
a ec ing
high
le els
o
ba
ac i i y
esul ed in
equa ions
wi h
high
p edic i e
powe
and
o
pa icula
alue
o
o e-
cas ing
he
e ec s
o
changes
in
land
use
on
ba s.
Al hough
espe ilionids
use
a
di e si y
o
habi a s,
he
eg ession
models
iden i y
ipa ian
and
woodland
habi a s
as
pa icu-
la ly
impo an .
Once
mo e
nume ous
and
widesp ead,
woodland
habi a s
a e
now
pa chily
dis ibu ed
and
u he
habi a
agmen a ion
may
educe
he
alue
o
hose
ha
emain
(B igh
1993).
Thus,
o
conse a ion
pu poses
he
ela i e
magni ude
o
such
an
e ec
may
be
p edic ed
using
da a
collec ed
in
he
su ey.
The
p incipal
ca ea
in
in e p e ing
he
esul s
o
he
su ey
is
ha
hey
e lec
he
habi a
p e e ences
o
he
pipis elle,
he
mos
abundan
ba in
he
Uni ed
Kingdom
(Ha is
e
al.
1995).
In
u u e
U.K.
su eys,
i
is
hoped
ha
olun ee s
will
be
be e
able
o
iden i y
ba s
o
species
o
species
g oups.
To
achie e
his
goal,
aining
cou ses
in
he
use
o
ba
de ec o s
a e
being
held
h oughou
he
coun y
Conse a ion
Ecology
o
Eu opean
Ba s
251
Table
17.1
Habi a
Types
Selec ed,
Used
in
P opo ion
o
A ailabili y,
and
A oided
by
Ba s
in
B i ain
GOOD
HABITATS
"<
Selec ed
in
all
land
classes
T eeline
B oad-lea ed
woodland
edge
Lake
o
ese oi
-•
POOR
HABITATS
Selec ed
in
some
land
classes;
ne e
a oided
Hedge ow
S eam
Coni e ous
woodland
edge
Mixed
woodland
edge
B oad-lea ed
woodland
opening
Mixed
woodland
opening
Felled
woodland
Ri e
o
canal
Pond
Selec ed
in
some
land
classes
and
a oided
in
o he s
Open
di ch
Co e ed
di ch
S one
wall
Coni e ous
woodland
opening
Sc ub
Pa k
land
U ban
land
A oided
in
some
land
classes;
ne e
selec ed
Imp o ed
g assland
Semi-imp o ed
g assland
Lowland
unimp o ed
g assland
A oided
in
all
land
classes
A able
Moo land
Upland
unimp o ed
g assland
No es:
The
esul s
in e p e ed
he e
a e
om
Walsh
and
Ha is
(1996a,
1996b).
"Land
classes"
e e s
o
he
19
disc e e
land
classes
in
hei
s udy.
(Ca o
1994),
ollowing
a
success ul
ini ia i e
in
he
Ne he -
lands.
The
esul s
o
a
na ionwide
ba
dis ibu ion
su ey
in
ha
coun y,
also
in ol ing
olun ee s
(Kap eyn
1991),
ha e
been
ecen ly
published
(Limpens
e
al.
1997).
A
de ec o
su ey
o
ba dis ibu ion
also
has
been
comple ed
in
Den-
ma k
(Baagoe,
in
manusc ip ).
Landscape
Ecology:
F om
he
Na ional
o
he
Local
Scale
In
a
majo
s udy
o
habi a
use
in
he
p o ince
o
Uppland,
Sweden
(59°
N),
de
Jong
(1994)
ound
ha
ela i ely
open
deciduous
o es s
and
adjacen
shallow
eu ophic
lakes
we e
he
only
habi a s
a ac ing
la ge
numbe s
and
a
high
di e si y
o
ba s
du ing
ea ly
summe ,
as
a
esul
o
high
chi onomid
p oduc i i y
(de
Jong
and
Ahlen
1991).
Ba s
o aged
in
mo e
di e se
habi a s
la e
in
summe ,
al hough
lakes,
we lands,
and
deciduous
o es
emained
impo an .
In
a
landscape
mosaic,
pa ch
size
is
impo an ,
and
he
numbe
o
species
o aging
in
a
pa ch
o
a o able
habi a
signi icandy
dec eased
when
he
a ea
was
less
han
30
ha
and
was
mainly
a ec ed
by
he
abundance
o
deciduous
o es
in
he
pa ch
and
by
he
ex en
o
i s
isola ion
om
o he
pa ches.
Species
ha
a oid
open
a eas
(Myo is
spp.,
Pleco us
au i-
us)
a e
less
likely
o
use
such
pa ches,
bu
in
some
cases
pa ches
a e
only
used
by
Nyc alus
noc ula.
The
composi ion
o
he
mosaic
also
a ec s
species
numbe ,
and
ewe
spe-
cies
we e
ound
in
pa ches
isola ed
by
open
ields
and
clea -
cu s.
Dense
closed-canopy
o es
was
a oided
by
all
species.
Obse ed
p e e ences
we e
o
ela i ely
open
coni e ous
o es
by
Myo is
b and i
and
M.
na e e i,
o
deciduous
o -
es
by
Pipis ellus
pipis ellus,
and
o
o es
edge
and
glades
by
P.
pipis ellus
and
Ep esicus
nilssoni.
Co ido s
o
ees
ha
connec
eeding
pa ches
a e
impo an ,
pa icula ly
o
M.
na e e i.
The
co ido
concep
has
an
in ui i e
appeal
and
has
been
widely
adop ed
by
ecologis s
and
land-use
planne s
in
ad ance
o
o mal
p oo
o
i s
alidi y.
Recen
e iews
(Hill
e
al.
1993;
Spelle be g
and
Gaywood
1993;
Dawson
1994)
concluded
ha ,
wi h
ew
excep ions,
ele an
s udies
lacked
s a is ical
igo .
The
use
o
linea
landscape
ele-
men s
by
ba s
has
been
demons a ed
in
he
Ne he lands,
whe e
in
he
open
landscape,
lanes,
a enues
o
ees,
hedge ows,
and
canals
a e
used
as
ligh
pa hs
(Limpens
e
al.
1989;
Limpens
and
Kap eyn
1991).
In
a eas
whe e
linea
landscape
elemen s
a e
abundan ,
ba
de ec o
su -
eys
e ealed
a
dense
ne wo k
o
ligh
pa hs
and
o aging
a eas.
In
mo e
open
a eas
de oid
o
such
elemen s,
ew
ligh
pa hs
and
ba s
a e
eco ded.
Some
species,
such
as
Myo is
dauben onii,
make
de ou s
along
hedge ows
a he
han
c oss
an
open
a ea
on
he
sho es
ou e
o
a
eeding
habi a .
In
compa ing
ba s
o aging
in
a able
land
and
in
an
adja-
cen
na u e
ese e
wi h
abundan
woodland,
Gaisle
and
Kolibac
(1992)
no ed ha
he
densi y
o
o aging
ba s
was
lowe
in
he
a mland
han
in
he
ese e,
by
an
o de
o
magni ude
o
Nyc alus
noc ula
and
by
a
ac o
o
wo
o
he
emaining
species.
In
a mland,
he
ba s
o en
lew
along
windb eaks,
which
a ac
and
p o ide
shel e
o
insec s,
and
ba s
equen ly
a el
on
hei
leewa d
sides
(Racey
and
Swi
1985;
Limpens
and
Kap eyn
1991).
Wind-
b eaks
may
also
educe
he
isk
o
p eda ion
(Scho ield
1996)
and may
be
used
by
ba s
o
o ien a ion
by
echoloca-
252
P.
A.
RACEY
ion
(Limpens
and
Kap eyn
1991).
Pipis ellus
na husii
is
hus
obse ed
mig a ing
sou hwes
in
au umn
along
Du ch
polde s
(P.
H.
C.
Lina,
pe sonal
communica ion).
In
a
i e
alley
mosaic
o
woodland
and
pas u e,
Racey
and
Swi
(1985)
showed
ha
P.
pipis ellus
used
a
egula
nighdy
ou e
and
lew
di ec ly
be ween
o aging
a eas.
Pipis elle
ba s
oos ing
in
ba
boxes
we e
s udied
in
sou he nmos
Sweden
(57°
N)
o
eigh
consecu i e
b eed-
ing
seasons
in
wo
con as ing
si ua ions:
a
150-ha
pine
plan a ion
adjoining
a
lake
and
a
16-ha
pa k
domina ed
by
deciduous ees o
a ying
ages
bu
su ounded
by
in en-
si e
a ming
and
indus y
(Ge ell
and
Lundbe g
1993).
The
popula ion
in
he
la e
a ea
declined
o e
he
s udy
pe iod,
while
ha
in
he
o me
inc eased.
This
di e ence
is
no
a ibu able
o
mo emen
o
he
popula ions,
as
Lundbe g
and
Ge ell
(1986)
ha e
shown
high
oos
si e
ideli y
in
his
species.
Body
mass
indices
(log,,
mass/log,
o ea m3)
we e
used
o
es ima e
a
ese es,
and
hese
we e
signi ican ly
lowe
in
3 o
4
yea s
in
Sep embe
in
he
a ea
adjacen
o
in ensi e
a mland.
Al hough
he
le els
o
o ganochlo ine
esidues
and
cadmium
we e
also
highe
in
he
popula ion
li ing
close
o
a mland,
Ge ell
and
Lundbe g
(1993)
consid-
e ed
ha
he
p oxima e
cause
o
he
decline
in
he
pipi-
s elle
popula ion
in
he
a mland
and
indus ial
a ea
was
he
de e io a ion
in
eeding
condi ions
caused
by
d ainage
and
wa e
pollu ion,
which
esul
in
dec eased
aqua ic
in-
sec
popula ions.
Because
o
he
con inued
p e e ence
o
he
noc ule
ba
Nyc alus
noc ula
o
oos ing
in
ca i ies
in
deciduous ees
a he
han
in
houses
and
he
loss
o
ma u e
and
pos -
ma u e
deciduous
ees
om
managed
landscapes,
con-
ce ns
ha e
been
exp essed
abou
he
long- e m
su i al
o
his
species
in
wes e n
Eu ope
(Hu son
1993).
Nyc alus
noc-
ula
does,
howe e ,
make
use
o
o he
an h opogenic
ac-
o s,
and
wo
s udies
(C anb ook
and
Ba e
1965;
K on-
wi e
1988)
ha e
shown
ha
i
o ages
on
house
c icke s
(Ache a
domes ica)
o e
domes ic
e use
dump
si es.
Ba s
and
S ee
Lamps
The e
ha e
been
a
numbe
o
s udies
in
Eu ope
and
No h
Ame ica
o
ba s
o aging
a ound
s ee
lamps
(Rydell
and
Racey
1995).
The
bluish-whi e
ligh o
me cu y- apo
s ee
lamps,
which
emi
ul a iole
adia ion,
a ac s
in-
sec s
whose
densi y
can
be
de e mined
by
lash
pho og a-
phy.
In
con as ,
low-p essu e
sodium
lamps,
which
emi
monoch oma ic
o ange
ligh ,
do
no
a ac
insec s,
and
high-p essu e
sodium
lamps
ha
include
some
me cu y
apo
and
hence emi
some
ul a iole
adia ion
a e
in e -
media e
in
e ms
o
insec
a ac ion.
Rydell
(1992),
using
a
ba
de ec o
ixed
o
a
mo ing
ca
(Ahlen
1980-1981;
Jiides
1989),
de ec ed
no he n
ba s
(Ep esicus
nilssoni)
a
ela i ely
high
densi ies
(2-5/km)
in
sou he n
Sweden
nea
whi e
s ee
lamps,
compa ed
wi h
0.1-0.4
ba s
pe
kilome e
o
unli
oad.
Means
o
3.2
and
3.1
pipis elle
ba s
(Pipis ellus
pipis ellus)
we e
eco ded
pe
kilome e
o
li
oad
in
En-
gland
and
Sco land,
espec i ely
(Blake
e
al.
1994;
Rydell
e
al.,
unpublished
da a).
The
g oss
ene gy
in ake
o
E.
nilssoni
o aging
a ound
s ee
lamps
was
mo e
han
wice
as
high
(0.5
kj/min)
as
ha
eco ded
in
woodland
(0.2
kj/min)
and
compa able
o
ha
o e
pas u e
whe e
he
ba s
o aged
on
dung
bee les
(0.6
kj/min).
S ee
lamps
may
allow
some ba
species
o
inc ease
hei
ene gy
in ake
and may
accoun
o
he
equen
occu ence
o
hese
species
in
buil -up
u ban
a eas
(Rydell
1992).
A
adio acking
s udy
showed
ha
noc ules
(Nyc alus
noc ula)
spen
mos
o
hei
o aging
ime
(~65%)
ei he
o e
a
lake
o
in
a
own,
hun ing
o e
an
asphal
su ace
(a
ca
pa k
and
oad
junc ion)
illumina ed
by
s ong
ligh s
(K onwi e
1988).
Adjacen
woods
and
a mland
we e
used
only
occasionally.
Typically,
he
ba s
ed
o e
he
lake
a
dusk
and
la e ,
a e
i
was
ully
da k,
mo ed
on
o
eed
o e
he
ligh s
in
he
own. Du ing
his
second
pe iod,
75%
o
he
o aging
ime
was
spen
o e
he
ligh s.
Simila
obse -
a ions
o
noc ules
in
England
eeding
o e
a
well-li
ail-
way
ya d,
which
subsequen ly
became
an
equally
well-li
p ison,
ha e
been
made
by
A.
J.
Mi chell-Jones
(pe sonal
communica ion).
In
a
con as ing
s udy,
Rachwald
(1992)
used
ba
de ec o s
o
moni o
noc ules
in
Bialowieza
p ime al
o es
in
Poland
whe e
he e
a e
se e al
illages
bu
no
s ee
lamps.
Ba
ac i i y
was
consis en ly highes
o e
wa e
bu
small
o es
clea ings,
main ained
o
adi ional
a ming,
we e
also
ex-
ploi ed.
Ba s
did
no
p e e
he
illages
o
o he
open
a eas.
In
a
adio acking
s udy
o
se o ines
(Ep esicus
se o inus)
in
England,
Ca o
e
al.
(1995)
showed
ha
second
o
ca le
pas u es
ha
p o ided
an
abundan
sou ce
o
Aphodius
dung
bee les,
oads
wi h
whi e
s ee
lamps
we e
he
mos
e-
quendy
used
eeding
si es.
In
sou he n
Swi ze land,
Ha ne
and
S u z
(1985-1986)
moni o ed
he
ac i i y
o
Pipis ellus
kuhlii
and
P.
pipis ellus
o
3
yea s
o e
abou
500
km
o
oad
using
a
ba
de ec o
on
he
oo
o
a
mo ing
ca ;
94%
o
P.
kuhlii
and
45%
o
P.
pipis ellus
we e
obse ed
nea
s ee
lamps.
The
equen
use
o
he
a ea
a ound
s ee
lamps
by
o aging
ba s
has
ob ious
implica ions
o
conse a ion.
In
con as
o
many
o he
ba
habi a s,
he
illumina ion
o
s ee s,
oads,
and
p i a e
p ope ies
is
inc easing
and
is
likely
o
bene i
a
leas
some
ba
species.
Po en ially
less
bene icial
has
been
a
ecen
endency
o
eplace
me cu y
apo
lamps
wi h
sodium
lamps,
which
use
less
ene gy
(and
a
less
haza dous
elemen )
bu
a e
o
less
alue
o
ba s.
Conse a ion
Ecology
o
Eu opean
Ba s
253
The
u ban
a eas
o
la ge
ci ies
ha e
deple ed
insec
aunas
(F ankie
and
Ehle
1978;
Taylo
e
al.
1978),
and
some
s udies
in
No h
Ame ica
ha e
shown
ha
ew
ba s
can
su i e
in
his
kind
o
habi a
(Geggie
and
Fen on
1987;
Ku a
and
Te amino
1992).
Howe e ,
a
majo
su ey
o
ba s
in
he
London
a ea
in
1985-1986
e ealed
137 ac i e
summe
oos s,
75
o
which
we e
pipis elles
(Pipis ellus
pipis ellus).
A
o al
o
430
summe
oos s
was
eco ded
du ing
a
36-yea
pe iod;
450
possible
eeding
si es
we e
su eyed
and
ba s
we e
eco ded
a
397
o
hese.
Pipi-
s elles
we e
he
mos
abundan
species,
al hough
Nyc alus
noc ula
and
N.
leisle i
we e
also
widesp ead.
In
con as ,
only
28
o
106
possible
hibe nacula
su eyed
we e used
by
ba s,
and
in
70%
o
hese
ewe
han
six
indi iduals
we e
ound
(Micklebu gh
1987).
Gaisle
(1979)
showed
ha
he
numbe
o
o aging
P.
pipis ellus
inc eased
om
he
sub-
u bs
o
he
ci y
cen e
o
B no
in
he
Czech
Republic
and
e iewed
he
common
occu ence
o
his
species
in
se e al
Eu opean
ci ies,
which
he
a ibu ed
o
he
p esence
o
s ee
igh s.
Di e en
species
o
ba s
a e
no
likely
o
be
equally
a ec ed
by
he
p esence
o
s ee
lamps.
Those
mos
likely
o
bene i
a e
ae ial
hawking
species
such
as
Nyc alus,
Ves-
pe ilio,
Ep esicus,
and
Pipis ellus.
Wi h
he
excep ion
o
Nyc alus
noc ula,
he e
is
li le
clea
e idence
ha
any
o
he
species
in
hese
gene a
is
p esen ly
h ea ened (Hu son
1993).
On he
o he
hand,
se e al
Eu opean
species
o
Myo is,
Pleco us,
and
Rhinolophus,
which
do
no
ake
ad an-
age
o
s ee
lamps,
ha e
su e ed
popula ion
declines
and
a e
endange ed,
a
leas
in
some
coun ies
(S ebbings
1988).
A
possible
nega i e
e ec
o
s ee
igh s
is
ha
hey
may
a ac
mo hs
ha
hen
become
una ailable
o
ba
species
which
a e
adap ed
o
gleaning.
The
echoloca ion
pulses
o
mos
Myo is
and
Pleco us
spe-
cies
lack
he
na owband
componen
necessa y
o
he
long- ange
de ec ion
o
insec s
in
open
ai
(Neuweile
1989).
I
may
be
ha
hese
ba s
a e
less
e icien
a
exploi -
ing
insec s
in
open
si ua ions
han
in
clu e ed
en i on-
men s.
Al e na i ely,
he
p eda ion
isk
in
b igh ly
ligh ed
condi ions
in
combina ion
wi h
open
si ua ions
may
be
oo
high
o
slow- lying
ba s.
E idence
om
high
la i udes
in
Scandina ia
and
Finland sugges s
ha
ba s,
pa icula ly
Myo is
spp.,
end
o
a oid
open
habi a s such
as
lakes
in
he
ambien
igh
condi ions
p e ailing
a ound
midsumme ,
bu
e u n
o
such
a eas
as
he
nigh s
become
da ke
la e
in
he
yea
(Nyholm
1965;
Rydell
1992).
Al hough
hese
indings
sugges
ha
he
mo emen s
o
Myo is
spp.
may
be
es ic ed
by
illumina ed
mo o ways
and
o he
i
a eas,
K ull
e
al.
(1991)
ound
ha
adio agged
M.
ema gina us
made
de ou s
along
a
mo o way
o
each
unde passes
lead-
ing
o
o aging
a eas.
Na ional
Roos
Su eys
The
i s
na ional
s anda dized
su ey
o
ba
oos s
in
En-
gland—The
Na ional
Ba s
in
Chu ches
Su ey—was
o ga-
nized
by
he
Ba
Conse a ion
T us
om
1992
o
1994
and
in ol ed isi s
o
538
chu ches
and
chapels
(Sa gen
1995).
Visi s
showed
ha
90%
o
132
chu ches
used
as
oos s
in
he
la e
1960s
we e
s ill
occupied
by
ba s.
The
mos
im-
po an
ac o
in luencing
he
likelihood
o
ba
occupancy
was
age
o
he
building,
bu
when
he
ela i e
e ec s
o
all
he
eco ded
ac o s
we e
aken
in o
accoun ,
including
aspec s
o
he
building
ela ed
o
age,
he
la e
dec eased
in
i s
le el
o
impo ance,
and
a
speci ic
combina ion
o
o he
ac o s
desc ibed
mo e
a ia ion
in
he
da a
han
age
alone.
The
ou
ac o s
ha ing
a
signi ican
e ec
on
he
p es-
ence
o
ba s
in
chu ches
and
chapels
we e
oo
ype,
wall
ma e ial,
geog aphic
loca ion,
and
he
le el
o
building
de elopmen
a ound
he
chu ch.
Ba s
we e
mo e
likely
o
occupy
chu ches
wi h
limes one
walls
and
lead
oo ing,
ea u es
ha
p obably
indica e
o he
p ope ies
which
a -
ac s
ba s
such
as
empe a u e
and
oos
si es.
In
gene al
ba s
occupied
chu ches
and
chapels
su ounded
by
p e-1800
buildings
and
a oided
hose
wi h
mode n
neighbo ing
buildings.
Isola ed
u al
and
illage
chu ches
we e
also
a-
o ed
by
ba s
a he
han
hose
in
u ban
a eas.
Au ecological
S udies
The
G ea e
Ho seshoe
Ba
The
g ea e
ho seshoe
ba
(Rhinolophus
e umequinum)
is
ca ego ized
as
an
endange ed
species
h oughou
much
o
Eu ope,
wi h
he
cu en
B i ish
popula ion
o
his
species
es ima ed
a
abou
4,000
indi iduals
di ided
among
abou
14
majo
ma e ni y
colonies
(Mi chell-Jones
1995).
Radio-
acking
s udies
ha e
e ealed
ha
ba s
usually
o age
wi hin
4
km
o
he
oos ,
and
he
conse a ion
o
o aging
habi a s
wi hin
his
adius
is
he e o e
pa icula ly
impo -
an
(Jones
and
Mo on
1992;
Jones
e
al.
1995).
Du ing
sp ing,
ba s
o age
in
ancien
semina u al
woodland,
bu
du ing
la e
summe
hey
eed
mainly
o e
pas u e.
The
ambien
empe a u e
in woodland
is
gene ally
highe
han
ha
o e
pas u e,
and
his
di e en ial
widens
as
empe a-
u e
dec eases.
Because
insec
abundance
inc eases
apidly
abo e
6°-10°C,
i
is
likely
o
be
mo e
p o i able
o
ba s
o
o age
in
woodland
in
sp ing.
The
shi
o
pas u es
du ing
he
summe
is
associa ed
wi h
he
dominance
o
Aphodius
dung
bee les
in
he
die
as
ca le
dung
accumula es.
The
abundance
o
such
bee les
may
be
h ea ened
by
he
use
o
A e mec in
an ihelmin-
254
P.
A.
RACEY
hics
in
ca le,
which
because
o
hei
pe sis ence
educe
he
insec
auna
associa ed
wi h
dung
(S ong
1992).
Because
ju enile
ba s
o age
independen ly
o
hei
mo he s
bo h
be o e
and
a e
weaning,
p ime
o aging
habi a
(ca le-
g azed
pe manen
pas u e
close
o
ancien
woodland)
adja-
cen
o
he
ma e ni y
oos
is
likely
o
be
impo an
o
ju e-
nile
su i al.
The
use
o
A e mec in
in
ca le
pas u ed
in
he
icini y
o
such
oos s
could
be
pa icula ly
de imen al
o
ju enile
ba s
(Du e ge
and
Jones
1994).
A
simila ly
de ailed
pic u e
o
he
cha ac e is ics
o
he
hibe nacula
o
g ea e
ho seshoe
ba s
has
been
assembled
by
Ransome
(1968,
1971,
1990).
These
hibe nacula
con ain
comple ely
da k
a eas
wi h
a
ela i e
humidi y
in
excess
o
95%,
a
ange
o
ambien
empe a u es
be ween
5°
and
10°C,
and
egions
o
slow
ai
low
esul ing
om
wo
o
mo e
en ances
o
a
sloping
en ance.
Close
access
o
shel e ed,
o en
sou h- acing
win e o aging
a eas
is
also
impo an ,
as
is
eedom om
epea ed
human
dis u bance.
As
a
esul o
he
p o ec ion
o
ma e ni y
oos s
and
hibe nacula,
he
decline
in
numbe s
o
g ea e
ho seshoe
ba s
(S ebbings
and
A nold
1987;
S ebbings
1988)
has
been
hal ed.
The
way
in
which
knowledge
o
he
au ecology
o
ho seshoe
ba s
is
being
applied
o
hei
conse a ion
in
England
and
Wales
has
been
de ailed
by
Mi chell-Jones
(1995)
and
includes
designa ing
key
oos s
as
si es
o
Special
Scien i ic
In e es
managed
by
he
ele an
s a u o y
con-
se a ion
agencies
(English
Na u e
and
The
Coun yside
Council
o
Wales).
The
ambien
empe a u e
o
ma e ni y
oos s
is
inc eased
by
he
use
o
hea e s
(Mi chell-Jones
1995),
and
ga es
ha e
been
ins alled
a
he
en ances
o
hibe nacula
o
p e en
dis u bance
(Ransome
1990).
A en-
ion
is
p esen ly
being
ocused
on
he
ways
in which
key
eeding
si es
a ound
ma e ni y
oos s
can
be
p o ec ed
(Mi chell-Jones
1995).
The
Lesse
Ho seshoe
Ba
His o ically,
he
lesse
ho seshoe
ba
(Rhinolophus
hip-
poside os)
oos ed
all
yea
a ound
in
ca es
(Ho acek
1984),
and
he
mos
ma ked
change
in
i s
oos ing
beha io
has
been
he
adop ion
o
buildings
as
summe
oos s.
In
a
su ey
o
he
cha ac e is ics
o
156
b eeding
oos s
in
he
Uni ed
Kingdom
(Scho ield
1996),
R.
hipposide os
selec ed
p edominan ly
nine een h-cen u y
buildings
(77%;
n
=
61)
wi h
s one
walls
(81%;
n
=
82)
and
sla e
oo s
(88%;
n
=
77).
Thei
oos ing
si es
we e
usually
loca ed
in
oo
spaces
o a ic
ooms
(95%;
n
=
76).
Access
o
oos s
was
com-
monly
h ough
la ge
openings
(>0.5
m2),
such
as
he
open
doo ways
o
windows
ha
o en
cha ac e ize
de elic
o
semide elic
buildings.
The
olume
o
b eeding
oos s
was
equen ly
g ea e
han
250
m3.
Simila
indings
a e
e-
po ed
om
o he
coun ies
in
Eu ope
(Gaisle
1963;
S u z
and
Ha lhe
1984;
McAney
and
Fai ley
1989).
Compa isons
o
a ailabili y
be ween
19
oos
buildings
and
20
andomly
selec ed
buildings
in
he
same
geog aphic
a ea
indica e
ha
buildings
used
as
oos s
a e
loca ed
clos-
es
o
s ands
o
b oadlea
and
mixed
woodland.
Buildings
con aining
oos s
a e
mo e
likely
o
be
connec ed
o
o ag-
ing
a eas
by
con inuous
linea
landscape
ea u es
such
as
hedge ows
o
s ands
o
ees.
A
Bon e oni
pai wise
compa ison
(Bye s
e
al.
1984)
be ween
he
land
classes
es ablished
(o
iden i ied)
by
he
Ins i u e
o
Te es ial
Ecology
(ITE)
(Bunce
and
Howa d
1991)
in
1-km
squa es
con aining
R.
hipposide os
oos s
and
hose
o
England and
Wales
as
a
whole
indica e
ha
his
species
selec s
o
hose
land
classes
associa ed
wi h
a eas
o
gen le
olling
and
undula ing
coun yside,
o en
enclosed
by
hedge ows
and
ee
lines. Land
classes
associa ed
wi h
la
open
coun yside
wi h
in ensi e
ag icul u e
o
exposed
upland
a eas
a e
a oided.
De ails
o
land
co e
(as
dis inc
om
land
class)
ob ained
om
sa elli e
da a
showed
ha
R.
hipposide os
selec s
deciduous
woodland,
and
ba s
we e
ound
o aging
as
much
as
2
km
om
b eeding
oos s
in
s ands
o
b oadlea
and
mixed
woodland,
ipa ian ees,
and
hedge ows.
Ba s
commu ed
o
and
om
o aging
a eas
and
hei
oos s
(bo h
b eeding
and
nigh
oos s)
along
con inuous
linea
landscape
ea u es
(pa icula ly
well-g own hedge-
ows).
In
addi ion
o
p o iding
o aging
a eas,
he
use
o
hese
ea u es
may
educe
p eda ion.
The
ambien
ligh
le el
and
heigh
a
which
ba s
lew
ac oss
a
5-m
gap
in
one
o
he
hedge ows
we e
eco ded
using
an
in a ed
ideo
cam-
e a.
A
ambien
igh
le els
g ea e
han
1
lux,
ba s
lew
a
a
heigh
o
less
han
1
m
om
he
g ound:
a
less
han
1
lux,
he
heigh
o
ligh
inc eased
o
4
m,
sugges ing
a oidance
o
a ian
p eda o s.
Conse a ion
s a egies
and
eco e y
p og ams
o
R.
hipposide os
mus
ake
in o
accoun
bo h
he
oos ing
and landscape
equi emen s
o
his
species.
The
B own
Long-Ea ed
Ba
BUILDING
ROOSTS.
In
a
s udy
o
he
oos ing
ecology
o
he
b own
long-ea ed
ba
(Pleco us
au i us)
in
no heas e n
Sco -
land
(57°
N),
En wis le
e
al.
(1997)
loca ed
56
oos s
in
buildings
and
compa ed
hei
cha ac e is ics
wi h
a
an-
domly
chosen
sample
o
buildings
om
he
same
a ea.
This
app oach
e ealed
ha
his
species
p e e en ially
oos ed
in
buildings
ha
we e
olde ,
highe ,
and
had
mo e
oo
com-
pa men s
(which
in
Sco land
a e
ully
ined
wi h
wood).
B own
long-ea ed
ba s
we e
ound
in
such
oos s
be ween
May
and
Oc obe ,
and
he
ypical
g oup
size
wi hin
he
oo
space
was
15-20
ba s.
Conse a ion
Ecology
o
Eu opean
Ba s
255
The
mean
empe a u e
wi hin
oos s
was
17.9°C,
and
oos s
we e
signi ican ly
wa me
han
a
andom
sample
o
buildings.
When
cap u ed
in
hei
oos s,
ba s
we e
gene -
ally
ac i e,
and
wa me
oos
empe a u es
may
ha e e-
duced
hei
dependence
on
o po .
The
empe a u e
inside
he
oos
was
posi i ely
co ela ed
o
he
equency
o
oc-
cupancy
and
also
wi h
he
size
o
he
ba 's
o ea ms,
wi h
la ge
indi iduals
being
cap u ed
in
wa me
oos s.
The
buildings
used
as
oos s
we e
close
o
ees
and
wa e
and
had
a
la ge
a ea
o
woodland
wi hin
0.5
km
han
a
andom
sample
o
buildings.
B own
long-ea ed
ba s
a e
oliage
gleane s
(Ande son
and
Racey
1991,
1993),
and
a-
dio acking
e ealed
ha
ba s
o aged
mainly
in
deciduous
woodland
in
he
icini y
o
he
oos ,
using
a
se ies
o
eeding
si es
o
which
hey
equen ly
e u ned
and
ha
we e
occasionally
sha ed
wi h
o he
ba s
om
he
same
oos s.
Females
spen
mos
o
hei
o aging
ime
wi hin
0.5
km
o
he
oos ,
whe eas
males
a eled
u he .
Ba s
e u ned
o
he
main
oos
on
77%
o
mo nings,
bu
also
used
al e na i e
si es,
which
had
coole
in e nal
empe a-
u es,
ollowing
colde
nigh s
(En wis le
e
al.
1996).
The
cha ac e is ics
o
he
oos
and
i s
loca ion we e
ela ed
o
he
ecology
and
beha io
o
i s
occupan s.
Ac oss
he
di e en
oos
si es
examined,
he
a ea
o
deciduous
woodland
wi hin
0.5
km
o
he
oos
was
co ela ed
wi h
o aging
pa e ns,
colony
size,
and
he
p og ess
o
he
male
ep oduc i e
cycle
(En wis le
1994).
Al hough
la ge-scale
dis ibu ion
pa e ns
o
his
species
a e
p obably
linked
o
a eas
o
woodland,
local
abundance
o
ba s
may
be
a ec ed
by
he
a ailabili y
o
sui able
oos
si es.
The
implica ions
o
he
wo k
o
conse a ion
and
managemen
a e
clea .
Building
oos s
a e
impo an
and
need
p o ec ion,
bu so
also
do
he
adjacen
woodlands,
which
should
be
main-
ained
and
hei
quali y
imp o ed
whe e
app op ia e.
Bu
i
building
oos s
o
b own
long-ea ed
ba s
ha e
a
sui e
o
cha ac e is ics
ha
dis inguish
hem
om
o he
oo
spaces,
why
a e
ba
boxes
a ac i e
o
his
species?
BAT
BOXES.
Ba
boxes
ha e
been
widely
used
o
many
yea s
h oughou
Eu ope
(Mayle
1990),
and
hei
occupancy
a e
is
highes
when
hey
a e
used
in
la ge
numbe s
in
coni e ous
plan a ions
de oid
o
na u al
oos
si es,
al hough
bi d
boxes
a e
also
used
by
ba s
in
deciduous
o es s
(Schlapp
1990).
In
he
1970s,
a
scheme
in ol ing
3,000
ba
boxes
was
ini ia ed
a
six
si es
ac oss
a
no h-sou h
ansec
in
he
Uni ed
Kingdom by
R.
E.
S ebbings.
F om
da a
collec ed
a
one
o
hese
si es,
The o d
Fo es ,
No olk
(53°
N),
one
o
he
ew
comp ehensi e
s udies
o
he
con ibu ion
o
such
boxes
o
he
popula ion
ecology
o
ba s
was
conduc ed.
Boyd
and
S ebbings
(1989)
analyzed
he
b own
long-ea ed
ba
occupancy
o
480
wooden
boxes
a ached
o
100
ees
a ound
he pe ime e
o
a
7-ha
ec angle
in
he
cen e
o
a
plan a ion
o
ma u e
Co sican
(Pinus
nig a
A nold)
and
Sco s
pine
(Pinus
syl es is
Linn).
The
boxes
we e
checked
wo
o
ou
imes
a
yea
o
10
yea s,
and
a
o al
o
219
emales
and
182
males
we e
cap u ed
and
indi idually
ma ked.
Following
he
es ablishmen
o
a
popula ion
o
ba s
in
he
boxes,
immig a ion
p obably
accoun ed
o
a
small
p opo ion
o
he
o al
ec ui men ,
he
emainde
coming
om
ep oduc ion
wi hin
he
popula ion.
The
mean
numbe
o
young
bo n
pe
emale
pe
yea
was
0.55,
and
he
o al
popula ion
inc eased
du ing
he
s udy
om
74
o
140
ba s,
gi ing
a
doubling
ime
o
10
yea s.
The
annual
su i al
a e
was
0.78-0.86
o
emales,
depending
on
he
me hod
o
es ima ion
used,
and
0.60
o
males,
simila
o
a es
ound
in
popula ions
o
b own
long-ea ed
ba s
oos -
ing
in
houses
in
no heas
Sco land
(En wis le
1994)
and
on
he
sou h
coas
o
England
(S ebbings
1970a).
Benzal
(1991)
s udied
b own
long-ea ed
ba s
occupying
520
bi d
boxes
in
a
130-ha
Pinus
syl es is
o es
a
he
com-
pa a i ely
high
al i ude
o
1,400
m
in
cen al
Spain.
The
boxes
we e
widely
used
in
he
i s
summe
a e
ins alla-
ion;
a
o al
o
197
indi iduals
was
ound
in
3%
o
box
inspec ions
and
d oppings
we e
ound
in
an
addi ional
8%.
Al hough
he
ba s
a i ed
in
he
s udy
a ea
in
he
i s
hal
o
May
and
s ayed
un il
he
beginning
o
No embe ,
hey
le
he
boxes
a
he
end
o
May
o
o m
ma e ni y
colonies
in
small
ca es
and
a ics
and
e u ned
o
hem
in
mid-July
wi h
lying
young.
These
s udies
p o ide
ecological
da a
ha
suppo
he
p omo ion
o
ba
boxes
as
al e na i e
oos s
o
ba s,
pa icula ly
in
a eas
de oid
o
such
oos s,
al hough
he
Spanish
s udy
sugges s
ha
boxes
may
no
always
be
app op ia e
o
b eeding.
Long- e m
s udies
a e
now
needed
o
compa e
he
popu-
la ion
dynamics
o
ba s
occupying
di e en
ypes
o
boxes,
pa icula ly
hose
made
om
sawdus
and
cemen ,
which
a e
o en
p e e ed
o
hose
made
only
o
wood
(Taake
and
Hildenhagen
1989;
Mayle
1990).
Despi e
he
posi i e
con-
se a ion
alues
o
ba
boxes,
hey
should
no
be
consid-
e ed
as
a
subs i u e
o
hollow
ees,
he
loss
o
which
is
a
majo
h ea
o
less
synan h opic
species
such
as
Nyc alus
noc ula.
The
Se o ine
Ba
The
se o ine
ba
(Ep esicus
se o inus)
is
widely
dis ibu ed
in
mainland
Eu ope
e en
a
mo e
no he ly
la i udes
(Schobe
and
G immbe ge
1993)
whe e
i
is
associa ed
wi h
highly
exploi ed
landscapes
and
appea s
o
be
inc easing
i s
ange
(Baagoe
1986).
By
con as ,
in
B i ain
his
species
is
la gely
es ic ed
o
sou he n
England
(Hu son
1991).
Radio ack-
ing
s udies
ha e
e ealed
ha
se o ines
in
B i ain
gene ally

256
P.
A.
RACEY
o age
wi hin
3
km
o
hei
oos s,
using
as
many
as
i e
o aging
si es
a
nigh
in
a
wide
ange
o
habi a s
ha
in-
clude
chalk
g assland,
sc ub,
pas u e,
and
a eas
a ound
whi e
s ee
lamps.
Se o ines
loca ed
and
exploi ed
empo-
a y
eeding
si es
such
as
ecen ly
mown
g ass
om
which
summe
cha e s
(Amphimallon
sols i ialis)
eme ged
(Ca o
e
al.
1995).
The
inc easing
incidence
o
Aphodius
bee les
in
he
die
o
se o ines
as
summe
p og essed,
eaching
a
peak
o
85%
o
iden i iable
agmen s
in
he
eces
in
Augus ,
con i med
he
impo ance
o ca le
pas u e
as
eeding
habi-
a
o
his
species
(Ca o
e
al.
1994).
Ca o
e
al.
(1995)
concluded
ha
se o ines
a e
well
adap ed
o
an
an h opo-
genic
en i onmen .
They a e
s ongly
philopa ic
o
hei
oos s
in
houses
ha
a e
loca ed
close
o
a
ange
o
eeding
si es
whe e
hey
can
ake
ad an age
o
cu en
a ming
p ac ice
and
s ee
lamps.
Dauben oiTs
Ba
Dauben on's
ba
(Myo is
dauben onii)
o ages
almos
exclu-
si ely
o e
wa e
and
a ound
ipa ian
ege a ion
(Jones
and
Rayne
1988;
Kalko
and
Schni zle
1989),
eeding
oppo -
unis ically
on
insec s
ha
swa m
in
such
si ua ions,
p in-
cipally
chi onomids
(Swi
and
Racey
1983;
Koku ewicz
1995).
Inc eased
numbe s
o
his
species
o
ba ,
some imes
o
se e al
hund ed
pe cen ,
ha e
been
epo ed
om
coun s
o
hibe na ing
indi iduals
in
he
Ne he lands
(Daan
1980;
Vou e
e
al.
1980;
Wein ich
and
Oude
Voshaa
1992),
Czech
and
Slo ak
epublics
(Ba a
e
al.
1981;
Ce eny
and
Bu ge
1990),
and
sou hwes
Ge many
( on
Hel e sen
e
al.
1987).
Koku ewicz
(1995)
pos ula ed
ha
he
inc ease
in
Dau-
ben on's
ba
is
a ibu able
o
eu ophica ion
and
canaliza-
ion
o
wa e ways
h oughou
Eu ope,
ac o s
which
in-
c ease
he
numbe s
o
chi onomids.
This
hypo hesis
was
ecen ly
es ed
in
no heas
Sco land
by
compa ing
he
ac i i y
o
pipis elles
and
Dauben on's
ba s
in
wo
i e s
wi h
sha ply
con as ing
ni a e
le els,
one
o
which may
be
designa ed
as
a
ni a e-sensi i e
a ea
by
he
Eu opean
Union
(Racey
1998).
This
s udy
p o ides
some
suppo
o
Koku ewicz's
hypo hesis
and
is
one
o
he
ew
examples
o
a
bene icial
e ec
o
aqua ic
pollu ion
on
ba
popula ions.
Hidden
Biodi e si y
Many
o
he
ecen
addi ions
o
Eu opean
na ional
aunas
ha e
been
ba s.
Pleco us
aus iacus
was
i s
desc ibed
in
he
Uni ed
Kingdom
by
S ebbings
in
1967.
Al hough
Topal
(1958)
i s
ques ioned
he
s a us
o
subspecies
o
Myo is
mys acinus,
i
was
no
un il
mo e
han
a
decade
la e
ha
Hanak
(1971)
e en ually
dis inguished
Myo is
b and i
as
a
sepa a e
sympa ic
species.
These
wo
sibling
species
a e
now
widely
ecognized
h oughou
Eu ope
(Baagoe
1973;
Co be
and
Ha is
1991),
and
ecen
molecula
wo k
by
Neme h
and
on
Hel e sen
(1993)
has
poin ed
o
he
exis-
ence
o
a
hi d
Eu opean
species
in
he
M.
mys acinus
g oup.
Jones
and
an
Pa ijs
(1993)
desc ibed
wo
phonic
ypes
o
Pipis ellus
pipis ellus,
and
e idence
is
amassing
ha
hese
a e
sepa a e
species
(Ba a
e
al.
1995).
I
also
aises
he
ques ion
o
whe he
addi ional
c yp ic
ba
species
emain
o
be
ecognized.
Pipis ellus
na husii
is
a
mig a o y
species
(S elko
1969)
and
mo es
sou hwes
om
he
Bal ic
s a es
in o
he
low
coun ies
in
au umn.
The
i s
b eeding
colony
o
P.
na husii
was
ecen ly
eco ded
in
he
Ne he lands
(Kap eyn
and
Lina
1994),
and
since
i s
i s
appea ance
in
B i ain
in
1969
(S ebbings
1970b)
i
has
been
eco ded
he e
wi h
inc eas-
ing
equency
in
au umn
(Speakman
e
al.
1991b)
and
sum-
me
(Rydell
and
Swi
1995;
Ba low
and
Jones
1996).
Conclusions
The
i s
na ional
ba
habi a
su ey
o
a
Eu opean
coun y
concluded
ha
he
main
o aging habi a s
o
espe ili-
onids
a e
associa ed
wi h
b oad-lea ed
woodland
and
wa e .
Many
au ecological
s udies
indica e
ha
hese
con-
clusions
a e
widely
applicable
h oughou
Eu ope,
and
one
o
he
bene i s
o
he
Eu opean
Union
is
ha
ini ia i es
o
es o e
woodland
and
o
main ain
he
quali y
o
i e s
ap-
ply
in
all
membe
s a es.
Fo
example,
he
long
decline
in
o al
woodland
co e
in
he
Uni ed
Kingdom
con inued
un il
1920
when
woodland
occupied
only
5%
o
he
U.K.
land
su ace.
Since
hen,
a
la ge
e o es a ion
p og am
has
inc eased
o al
woodland
co e
o
10%.
Al hough
coni e s
accoun
o
mos
o
his
inc ease,
he
p opo ion
o
b oadlea
ees
plan ed
each
yea
has
inc eased
10- old
since
1985
and
now
exceeds
coni-
e
plan ing,
so
ha
a
hi d
o
he
2.4
X
106
ha
o
wood-
land
in
he
Uni ed
Kingdom
includes
b oadlea
ees.
This
change
has
been
la gely
d i en
by
he
p o ision
o
gene -
ous
inancial
incen i es
o
b oadlea
plan ing,
no
only
because
o
i s
aes he ic
alue
bu
because
o
a
wide
app e-
cia ion
o
he
alue
o
such
ees
o
many
aunal
g oups
(Osbo ne
and
K ebs
1981;
Kennedy
and
Sou hwood
1984).
Simila
g an
aid
has
been
a ailable
o
he
es o a ion
o
ponds,
75%
o
which
ha e
been
los
in
B i ain
since
1880
(Als op
and
Biggs
1993).
Such
schemes
ha e
he
added
alue
o
p o iding
good
o aging
habi a
o
ba s
and
coun-
e ing
he
end
owa d
habi a
agmen a ion.
Un o u-
na ely,
he
a e
o
loss
o
hedge ows, which
a e
impo an
landscape
elemen s
o ba s,
s ill
exceeds
he
a e
o
hei
eplacemen
(Ba
e
al.
1994).
The
widesp ead
use
o
ag icul u al
pes icides
and
eme-
Conse a ion
Ecology
o
Eu opean
Ba s
257
dial
imbe
ea men s
wi h
o ganochlo ides
may
ha e
had
majo
dele e ious
e ec s
on
ba
popula ions
in
he
1950s
and
1960s.
Res ic ions
on
he
mos
pe sis en
and
oxic
chemicals
may
ha e
esul ed
in
he
epo ed
inc ease
in
numbe s
o
some
ba
species,
simila
o
he
eco e y
ha
has
occu ed
in
ap o s.
Pollu ion
is
also
being
educed
in
many
i e s,
al hough
eu ophica ion
may
ha e
a
bene icial
e ec
o
hose
species
ha
eed
o e
wa e .
The
p incipal
way
in
which
he
alue
o
coni e
plan a-
ions
is
being
enhanced
o
ba s
is
by
he
p o ision
o
oos -
ing
boxes,
and
iable
and
sel -sus aining
popula ions
o
ba s
become
es ablished
in
such
boxes
when
la ge
numbe s
a e
p o ided
in
ela i ely
small
a eas.
Fu he
esea ch
is
needed
on
he
popula ion
dynamics
o
ba s
inhabi ing
boxes
made
o
sawdus
and
cemen ,
which
a e
mo e
a ac i e
o
hem
han
wooden
ones.
Recen
s udies
ha e
de ailed
wi h
some
s a is ical
igo
hose
physical
ea u es
associa ed
wi h
buildings
used
by
ba s
as
oos s.
Se e al
espe ilionid
and
hinolophid
species
o -
age
wi hin
a
ela i ely
sho
dis ance
o
such
oos s
so
ha
he
conse a ion
and
managemen
o
adjacen
woodland
is
pa icula ly
impo an .
One
o
he
mos
aluable
a mland
habi a s
o
ba s
is
unimp o ed
pas u e,
whe e
ba s
o age
on
insec s
wi h
sub e anean
la al
s ages,
such
as
cha e s,
o
on
dung
bee les.
Fa ming
p ac ices
such
as
he
use
o
an i-
helmin hics
ha
educe
he
insec
auna
associa ed
wi h
dung
o
which
educe
he
a ailabili y
o
dung
such
as
ze o-
g azing
( anspo ing
cu
g ass
o
ca le)
may
ad e sely
e -
ec
se e al
ba
species.
Ba s
gene ally
o age
on
a
a ie y
o
insec
g oups,
and
we
need
o
know
whe he
he
loss
o
ce ain
axa
can
be
accommoda ed
by
a
shi
owa d
o he s.
In
some
cases,
howe e ,
pa icula ly
o
highly
endange ed
species
such
as
ho seshoe
ba s,
inancial
incen i es
a e
needed
o
s imula e
managemen
o
a mlands
nea
oos s.
Ba s
a e
among
he
mos
synan h opic
mammals,
elying
on
human-made
s uc u es
o oos ing
in
bo h
u al
and
u ban
a eas.
They
eed
oppo unis ically
on
concen a ions
o
insec s,
such
as
hose
a ound
whi e
s ee
ligh s
and
domes ic
e use
dumps.
The
use
o
whi e
s ee
ligh s
may
ha e
some
conse a ion
alue
because
he
ene gy
in ake
o
such
ba s
is
enhanced.
Un o una ely,
hose
species
exploi -
ing
his
eeding
oppo uni y
a e
less
h ea ened
han
hose
ha
do
no .
In
many
Eu opean
coun ies,
ba s
a e
he
mos
impo -
an
con ibu o s
o
mammalian
biodi e si y,
and
he
use
o
ba
de ec o s
and
applica ion
o
he
powe ul
echniques
o
molecula
gene ics
con inues
o
add
ba
species
o
na ional
aunal
lis s.
This
is
po en ially
impo an
because
a
he
1992
Rio
Con en ion
on
Biological
Di e si y
many
pa ici-
pa ing
poli icians
pledged
o
hal
he
wo ldwide
loss
o
animal
and
plan
species
and
gene ic
esou ces.
The
i s
con e ence
o
he
pa ies
o
he
Eu opean
Ba s
Ag eemen
held
in
he
Uni ed
Kingdom
in
1995
ag eed
o
a
wide- anging
conse a ion
and
managemen
plan,
which
in ol es
su ey
and
moni o ing
o
popula ions,
he
iden i-
ica ion
and
p o ec ion
o
impo an
oos s
and
o aging
habi a s,
and
he
p omo ion
o
public
awa eness
abou
ba s.
The
majo i y
o
hese
goals
will
be
achie ed
by
a
s ong
olun a y
sec o
willing
and
able
o
unde ake
su eys
and
moni o ing,
unde pinned
by
he
wo k
o
ecologis s
in
iden-
i ying
and
cha ac e izing
impo an
oos s
and
o aging
habi a s.
Acknowledgmen s
I
am
g a e ul
o
he
ollowing
colleagues
who
commen ed
on
a ious
d a s
o
his
chap e :
H.
Baagoe,
J.
Bu on,
C.
M.
C.
Ca o,
M.
B.
Fen on,
J.
Gaisle ,
A.
M.
Hu son,
G.
Jones,
S.
Micklebu gh,
J.
Rydell,
G.
Sa gen ,
H.
W.
Scho ield,
and
A.
L.
Walsh.
I
hank
L.
Young
o
he
alian
e o s
a
he
keyboa d.
Li e a u e
Ci ed
Ahlen,
I.
1980-1981.
Field
iden i ica ion
o
ba s
and
su ey
me h-
ods
based
on
sounds.
Myo is
18-19:128-136.
Ahlen,
I.
1981.
Iden i ica ion
o
Scandina ian
ba s
by
hei
sounds.
Repo
6,
Depa men
o
Wildli e
Ecology,
The
Swedish Uni-
e si y
o
Ag icul u al
Sciences,
Uppsala.
Als op,
C.,
and
J.
Biggs.
1993.
P oceedings
o
he
Con e ence
on
P o ec ing
B i ain's
Ponds.
Wild owl
and
We land
T us
and
Pond
Ac ion,
Slimb idge,
U.K.
Ande son,
M.
B.,
and
P.
A.
Racey.
1991.
Feeding
beha iou
in
cap i e
b own
long-ea ed
ba s
Pleco us
au i us.
Animal
Beha -
iou
42:489-493.
Ande son,
M.
B.,
and
P.
A.
Racey.
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.
Animal
Beha iou
46:1151-1155.
Baagoe,
H.
J.
1973.
Taxonomy
o
wo
sibling
species
o
ba s
in
Scandina ia:
Myo is
mys acinus
and
Myo is
b and i
(Chi op e a).
Videnskabelige
Meddelelse
a
Dansk
Na u hs o isk
Fo ening
136:191-216.
Baagoe,
H.
J.
1986.
Summe
occu ence
o
Vespe ilio
mu inus
(Linne
1758)
and
Ep esicus
se o inus
(Sch ebe
1780)
(Chi op e a,
Mammalia)
on
Zealand,
Denma k,
based
on
eco ds
o
oos s
and
egis a ion
wi h
ba
de ec o s.
Annalen
de
Na u his o i-
schen
Museums
in
Wien
Se ie
B
88-89:281-291.
Ba low,
K
E.,
and
G.Jones.
1996.
Pipis ellus
na husii
(Chi op e a:
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o
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Ca es
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30:75-84.
18
Impac s
o
Igno ance
and
Human
and
Elephan
Popula ions
on
he
Conse a ion
o
Ba s
in
A ican
Woodlands
M.
BROCK
FENTON
AND
I.
L.
RAUTENBACH
The
E hiopian
zoogeog aphic
egion
(sub-Saha an
A ica)
co e s
an
a ea
o
23,426,000
km2,
o
which
less
han
10%
his o ically
was
ain o es
(Keas
1972).
Al hough
he
e-
gion
is
bes
known
o
i s
la ge
mammals,
he
Chi op e a
include
mo e
species
han
any
o he
o de
o
mammals.
App oxima ely
174
species
o
ba s
li e
mainly
in
he
E hio-
pian
egion,
showing
a
high
le el
o
endemism
(24
o
41
gene a)
wi h
he
anges
o
only
abou
6
species
(3%) ex-
ending
beyond his
a ea
(Hayman
and
Hill
1971;
Nowak
1994).
App oxima ely
24
o he
species
whose
anges
ex end
in o
A ica
occu
mainly
ou side
he
E hiopian
egion,
mak-
ing
a
o al
o
abou
198
species
o
ba s
known
om
A ica
(Hayman
and
Hill
1971).
The
pu pose
o
his
chap e
is
o
conside
he
conse -
a ion
challenges
posed
by
ba s
in
A ican
woodlands.
We
selec ed hese
habi a s
o
wo
easons:
Fi s ,
ou
collec i e
expe ience
is
ocused
he e,
and
second,
he
woodlands
domina e
he
zoogeog aphic
egion
(Figu e
18.1)
and
o en
o e lap
wi h
a eas
o
high
human
popula ion
densi y.
In
his
chap e ,
we
use
he
nomencla u e
o
ba s
p esen ed
in
Nowak
(1994).
The
Gene al
Si ua ion
The
human
popula ion
in
sub-Saha an
A ica
is
p ojec ed
o
inc ease
i e- old
in
a
li le
mo e
han
a
hund ed
yea s,
om
0.6
billion
in
1990
o
2.8
billion
by
2100
(Bongaa s
1994).
In
1989,
he
human
popula ion
densi y
in
A ica
anged
om
2
o
265
(mean
±
SD,
41.8
±51.5)
pe sons
pe
squa e
kilome e
(S ua
e
al.
1990).
The
po en ial
impac
o
he
expanding
human
popula ion
o
he
conse a ion
o
A ican
bio a
canno
be
o e s a ed.
Fo
example,
since
1900,
he
human
popula ion
in
Zimbabwe
has
inc eased
om
0.5
o
mo e
han
10
million
(Cumming
1991),
and
in
he
pas
40
yea s
he
a e
o
land
clea ance
in
he
Sebungwe
Dis ic
o
Zimbabwe
has
been
abou
4%
pe
annum
(D.
H.
M.
Cumming,
pe sonal
communica ion).
Changes
o
his
magni ude
a e
expec ed
o
ha e
p o ound
e ec s
on
habi-
a s
and
hus
pose
a
majo
h ea
o
he
su i al
o
he
A ican
auna
(Ma in
and
de
Meulenae
1988).
The
di ec
and
indi ec
e ec s
o
an
expanding
human
popula ion
a e
expec ed
o
be
he
p incipal
ac o
causing
he
ex inc ion
o
many
o ganisms,
including
ba s.
261