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Ecological, beha iou al, and he mal obse a ions o a pe iphe al popula ion
o Indiana ba s (Myo is sodalis)
Con e ence Pape · Janua y 1996
CITATIONS
10
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Ecological,
Beha iou ill,
and The mai
Obse a ions
o a Pe iphe al
Popul ion
o
Indiana Ba s
(
N[yo is
sodalis)
ALLEN KURTA, KIMBERL ' ,J
. WILLIAMS' AND ROBERT MI
ES
ABSTRACT
We s udied
oos - ee
selec ion,
in e - oos
mo emen s,
and body em-
pe a u -e
oI z3
adul
e nale
ald ju enile
Indiana
ba s o e
a h ee-yea
pe iod.
Ba s
selec ed
hei
loos
ee on he
basis
oI species
and diame e ,
u e
amoun o ex blia ing
ba k was
no a ac o .
These ba s
oos ed
exclusi ely
in g een
ash
(F u , i l s
pennsyl anica) i h
a mean diame e
o
4o.9
! .z
cm (se;
l
= z3). Unlike
Indiana
ba s
in he
cen e o[ he
ange,
hose
in Michigan
ne e
used
sil e
maple
(Ace
saccha inum),
ne ; oos ed
in a shaded
a ea,
and ne e
oos ed
in a heal hy,
li ing ee.
The ba s
chalged
oos
lees
e e y
2.9
days,
on a e age,
mo ing om o
78
m o an al e na e
;ee.
P egnan
emales
changed
oos s
mos o en'
and lac a ing
ba s,
he leas .
O e all,
oos -swi ching
was no sa is ac o ily
explained
by hypo heses
based
on p eda o
a oidance,
in e up ion o
pai-asi e
li e cycles,
acking
o ood esou ces,
o en i onmen al
condi-
ions.
The epheme al
na u e
o ba k oos s
may
be a ac o
in equen
oos -swi ching,
bu i has
no been
adequa ely
es ed.
The numbe
o
ees occupied
by he g oup each
yea a ied
F om
i e o 18, al hough
wo
o bu ees
each
yea
we e
used
mos hea ily.
Con a y o published
in o ma ion,
hese
ba s
we e
no ad e sely
a ec ed
by bod empe a u es
o[ 35-4o'C.
INTRODUCTION
Guiclelines
b nanagemen
o enciange ed
species
gene ally
a e de i ed
om s udies
pe o med
nea
he cen e
o an animal's
geog aphic
ange'
whe e popula iops
a e la ge
and s able
and, he e o e,
easie
o wo k wi h
han
hose
on he
pe iphe y.
Ne e heless,
pe iphe al
popula ions
a e
wo hy o a en ion.
l one
accep s
he classical
iew ha pe iphe al
PoPu-
la ions
a e
wagilg
il cons an
ba le
wi h ex inc ion,
hen
he declining
heal h
o an
ou lying
g ouP
may be
an ea ly
wa lling o po en ial
p ob-
le ns
l'o
he
n ain
popula ion.
Con e sely,
ecen
allalyses
sugges
ha
pe ip ie al
populi ions
a e
mo e
likely o su i e
han cen ally
loca ed
ones,
and ul in a ely,
pe .iphe al
l eas nay become
c i ical
e ugia
o a
sECTIoN l Roos l lS
Ecologlcal
and
Beha lou al
Obse a lons
spccics
(l.o nolino
an l
(ll i nncll
cql).
I
any
c c l , a P uclcn
) anagc
n cn plan
slioulcl
spcci icall
adcl css
pc iphc al,
as wcll
as
cen l,
popu-
la ions, [ cca sc cgc a i c
u cl
cli ni ic co di ions
o icn
di -c
ac oss
al
spccics' angc,
ancl
guidclincs li n ula ccl
{b
onc g oup
n)ay
no
l c
applicablc o l c
o hc
(c.c.,
Lo is
99.1
).
- hc
In lia'a
ba a. cs
ac .ss
' ch i - hc
cas c ..
U.i ecl
S a cs
(- l o ison
gBz).
l)is u .bancc
lu ing liibc na ion
und
in cn ional
n ocji-
ica ion
o . najo
hibc ac'la
lccl <l
sc ious
cleclincs
in p.pula ion
sizc
du ing
hc l96os
and
ca .ly 97os, ancl
consccluc l ,
hc species
was
decla cci
cndansc ccl
in 97j
(llich c
c
al. 993).
Ilibe na i g
indiana
bi s
. e now cll
p|o cc ccl,
[ L hc
spccics
con i cs
o cicclinc,
sugges ing
p oblcn s
in su n ne
as . cll
(Olal son
9.S7).
Du ing hc wa n-wea he
no hs,
hcsc
ba s o< s
ilin a ily
unde .
cx blia ing
ba k, in upland
woocilo s
ancl lipa ian
i> cs s.'lb
cla e, sumnlc
oos ing
habi s
l' a e
been
nos ex cnsi ely
s udicd
in Illinois
((ia cl c .e
al. 99 )
and
Missou i
(Calahan
993),
in i c
hca
o hc
spccics'
dis ibu ion.
ln he p cscn
lcPo , c
dcsc ibc
a n ul i yca
s cly
o a s mme Po1' -
ula ion
o Incliana
ba s, oos ing
only 5
kn l.o n
hc
no hc n
cdge o
hei
ange.
OL goals wc c
o p o icle
bascline, ecological,
and behi -
iou al da a,
applicablc
o hc lnanilscmen
o his
specics
in no he n
a eas, z ncl
o n i kc
con PaIisons i h
p e ious
s uciics pe [o med
in he
co e
o he spccies' angc.
wc e c
p.l cula|ly
c cs ecl
in ac o s
a ec
inc oos
selec ion
and oos
lideli y,
b l e
also
cxa nined
he ques ion
o
body en pc a - c.
Ilasccl
on lal ola o
cxpc i nen s,
Hensha
and Folk
(1966)
sugges ed
ha
body en pe a u es
o only 34-35oC
we e
h al o his
ba . I ue,
such a low,
le hal
body empe a u e
ould
ha e a p o ound
ccological in pac ,
sl,e ely li ni ing
he
ypes
o oos
si es ha his ee-
dl clling
ani al
co - lcl
usc.
METHODS
The s udy
ook place
nea Ve mon ille,
Ea on
Coun y, Michigan, o n
3o
June
99
o 5 Augus
99j.
Indiana
ba s we e
cap u ed
in mis ne s
hile
lying
o e s eams
o in ligh pa hs
leading
om hei
oos s. A e
cap u e,
son e
do sal u as
i n ned i h
scisso s,
and
a o.7-g adio-
ansn i e
(Holohil
Sys en s
Inc., woodlawn,
on a io) was
a ached
using
a su gical
adhesi e
(Skin-llond).
The oos
ee was
loca ed
on subsequen
clays, L sing
a h cc-clen cn
Y gi an cl na
and ecci e
(Wildli e
Ins u-
n cl s,
'RX-20oos).'l)' ns ni c s
cn ined
a achecl
and unc ional
o u;
o u d.lys.
I)opula ion
sizc
*'as
lc c inccl
by coun ing
ba s as hey le hei oos ,
be een
sunse nd
5o
n inu cs
alic sunse .
we consis en ly
obse ed
ces
ha
shel c ed an Incliana
bi
i h
a unc ioning
ansmi e . in addi ion,
c
nadc coun s
a o l)c
oos ccs
(o iginally
dc ec ed b1' adio-
acking), . hen
mul iplc
obsc e s e e
a ailable,
o when e
did no
ha e a -ans ii e
on an
ba .
'lb clla ilc c ize
l e
hal i , ,
c cs ablishecl
o. -ha, ci cula
plo s
su -
o , ncling
ei ch
oos ,
and iclen ied
e e y
ee > 10 cnl DIIH wi hin he
plo s. I1-plo s
lb adjaccn
'oos s o',,e lappecl,
e
clnly used da a
om he
o3
The mal Obse a lons
S a ls lcs
Ma e nl y Roos
Pa ame e s
li s loos l i
i s
clisco c c l
o n , i ain
inclcpc lcncc
o1- hc
nicas ccl clian c c a [ ci s hcigh
( nu ) an l
clc c inccl
hc
cach oos cc a cl hc p cl'c ccl cxi 1.loin
sing; clinon c c .
cli ssi iccl cach cc i cco li g o l c an oL ol'loosc [ . k
ha
si s a sys cn cco nc lccl
[ y Gi ldnc c al.
( q9 ).
A a ing
as
gi c
o ccs i h > z5'Z
k osc
l i k, n cc'liun
) <l()6
bu <
low was ) o(X l u < o(k .
cia a.
Wc
hcigh
o
Wc also
wi s
p escl I
o high
25(Z),
and
In his cpo , wc includc i li na ion
ob ainc l l.o n
i c
o{-cigh oos
.ecs
p c iously clcsc .i[ c l
in Ku a c al. (199].1
).Wc clisca clcd
cla a
on
l lcc lccs
( clos s
I ,
C, and ll) n cn ionccl in ha pape
bccause hosc
oos s
wc c
disc c c l
only b acciclcn i lly ol sc ing
ba s
lea ing e
ees
. clusl<.
'l
hc lcccn lcn on-s a ion lia
a sin ila spccies,
hc no he n
L a
(M. s p io olis
), also oos ecl occasionally
unclc ba k in his
we lanci
([]os c
993), ncccssi a cd
l a
wc e
-c alua c
hc cla a
sing
only
ccs
loca ccl [ y i c ually
a lio- acking Indiana
l a s.
ll.aclio- ansn i e s uscd
i 99z ancl 993 wc c cn pc . u c.sc si i e,
and
all< ed us o de e ine hc
su ace cmpe a u e
o ' hc ba while
i was
in
hc oos . Da a wc e o
aken in a sys en i ic mi nne in 1992,
bu in
99j, body c l pc a Ll c as
cco clcd
c e y
3 l
ninu es
bc wccn
sun ise
. ncl sul sc . Du ing
bo li yea s,
a lbicn en pe a u c in hc shi de
was
sin ul ancc' usly neasu ed
', i l.
a n c cu y
he monlc c .
lo a oid dis u ,
bancc, cn pc i L cs'" c c
cco clccl ncl c
l an oo n l ' u
he oos .
Al hough l e l ould
Iike
o ha e noni o ed
oos ambien
empe a u e a
he same i le as
body empe a u e,
his was
no p ac ical
because, as
clcsc ibed
la c ,
he
ba s l equcn ly
changed oos s
and uscd
many di -
c en
ees.
All n eans a e ollo cd
by * 6'.'. s anda d e o
anc.l
hcn he ange
in
pa en hescs.
Co npa isons
be een o g oups we e
pe o n ed
using he
nolmal app oxi la ion
o he
Wilcoxon
Two-sa nplc
Tcs , whe eas
es s
in ol ing n o .e han
wo ca ego ies elied
on he
chi-sclua ed
app oxima-
ion o he
K uskal-Wallis
Tes .
l'es s in ol ing
enume a ion
da a
(con-
ingency ables) we e
pc o med
using chi-squa ed
es s
o , when
expec ed
equencies
ell below
i e, Fishe 's
Exac Tes . We used
Spea man's
Rank
-l'cs
o examine po en ial
co ela ion
be ween a iables.
All es s we e
wo-
ailed and uscd
a signi icancc
le el o o.o5.
RESUTTS
We
adio- lacked
z3 lndiana
ba s and loca ed he oos
ce
on i3o o l4
clays
ha he ansn' i c s
e e
unc ional.
The ba s included
six p egnan
incli iduals,
se cn
li c a ing
ba s, i e pos -lac a ing/non- ep oduc i e
cn ales, and i e
ju enilcs.
l'hc ba s oos ed p ima ily
unde he ex olia -
ing
ba k o z3 di e enl
ces.
All oos
ees we e dead
o nea ly dead
g ccn ash ha hacl
los n os
o all o hci najo
b anches; any li ing
blanches
hi
e naincd
c e <l cln in dian e e .
A e age oni o oos
ees was
4o.9
! .z
(3o-52)
cn . Mean oos - ee
heigh
was
z5.
* .6
(B-+l) n, and
nean
cxi heigh was
9.9
o.9
( .a- 8)
m. Iloos
ees
s ECT o N Roos ng
Analysls
o WoodY
Vege a lon
E ldence
o Roos
Selec l l Y
Roos
Loyal y
Wl hln Yea s
we e on a c age
)8.7
+ /. ( +/) l away
l o n lno he cc
se..l
i s
a
oos
in hc
samc
yca ald V3 4.5
( - ) m away
om he nea es ee
used
as
a oos
du ing
any
o he
h ce
yea s.
The e
we e no signi ican
di e cnces
among
yea s
in ouH, heigh ,
exi
heigh ,
o nea es -neighbou
clis ance
(all p >o. z).
All oos s
we e
in an unshacled,
5-ha
we land and
su ounded
by up o
n o s anding
wii e .
Ou cge a ion
analysis
yielded
nine
non-o e lap-
ping
plo s
ha
con ailed
99
g een
ashes,
34
sil e
maples,
nine Ame ican
elms
( U/mus
an e icann),
and o
ba kless,
uniden i iable
unks.
Al hough
his a ea
was o es ecl
p.ls u e
a one ime' i has been
pe -
sis en ly
inunda ed
b 5 yca s,
leading
o he
dea h o mos ees
and he
i nminen
dea h
o he
es . Consecluen ly,
many
ees
aPPea ed sui able
o Indiana
ba s
(i.e.,
loose
and
peeling
ba k
was p esen
on he unks).
Wi hin ou plo s,66
g een
ashes
e e sui a le
(16
ac ual
oos s
and
50
appa en ly
unused),
as
wc e
z sil e
maples,
and wo Ame ican
elms'
Because
o he s nall
sample
o elms,
we excluded
hem om u he
analyses.
We e
Indiana
ba s
selec i e
in choice
o oos ee?
We examined
h ee
po en ial
modes
o selec i i y-by ee
species,
DBH,
and he amoun o
ex olia ing
ba k p esen
on he unk' Wi hin ou plo s,
760/o
o he 87
po en ial
oos
ees
(excluding
elm) we e
g een ashes,
whe eas
z o/o
we e
sil e
maples.
Using
hese
alues
as expec ed
p opo ions,
we de e mined
ha he p obabili y
o all z3 oos s
being
g een ash
as
he esul
o chance
was less
han o.oo5
(X2
= 7'32;
d' '= ), and
we concluded
ha
he ba s
u'e e choosing
g een
ash
o e
sil e
maple.
We e
Indiana
ba s
mo e likely o use
a ee
wi h a la ge amoun o
loose
and
peeling
ba k?
ollowing
Ga dne
e al' ( 99 ),
we anked
lz
oos
ees
as high, nine
as
medium,
and wo as low.
This dis ibu ion di -
e ed
om andom
(X2
= 6'87;
d' '= '; p <o'o )' and supe icially
sup-
po s
he hypo hesis.
Ilowe e ,
wi hin ou sample
o 5o sui able
g een
ashes
ha we e
no used
as
oos s,
he dis ibu ion
o high, medium, and
low was z , 13, and 16,
espec i ely'
The e
was no signi ican
di e ence
(X'= 4.73;
d. .= ,; p >o.o9) be ween
oos
and
non- oos
samples,
and
hence,
no e idence
ha he amoun o loose
ba k was a ac o
in oos
selec ion.
Al hough
he a noul o loose ba k was no impo an ,
he size
o he
ce
was. The clia e e
o( oos
ees
(mean
= 4o.g
* .z cm; n = 23)
was
signi ican ly
g ca e
(z = j.B5;
p <o.oool)
and signi ican ly
less
a iable
(1.-,,.
.,,
= j.03;
| = o.oo6)
han
he diame e
o sui able,
non- oos
g een
ashes
(n eiln
- 3l4
4 c n;'l = 5o)
wi hin ou
plo s.
Diame e
o non-
oos
ees
a ed
o n ll o 70
cm, whe eas
ees used
by lemales
and
young
we c igh l,
g oupcd
be ee
n 3o
and
52
cm'
Indi iduals
wc e no highly
ai hiul
o a pa icula
ee.
Se en een
oI he
2l ba s
s i c[c{
ecs
a leas o ce,
yielding
a o al o 37
oos
changes.
O
he day
l a a ba
as i s
loca cd,
e
did no know
whe he
i had
changccl
oos s o ha1|
cn ained
in hc Sanle
lcc; he e o e,
we dele ed
he i s clay
l- 'o o -
alalyses.
1-hi y
sc en
oos
changes
o e he
cl aining
o7 6bsc a ion
clays
inclica ecl
[a Indiana ba s
ypically
105
Ll .u llc l
()(). s
' ,c ),
.9 l. 's.
.lo l. l
nu l c
ol oosI
cl lic.s/ a
as
olll)' ,o,
b
chi ngcs/[ .
a iccl
l< n
zclo o six. Nul c
o -changcs
as
p<lsi i cly
c<l l'cla ccl
', i l
n[ c
o 'clays
l a
hc l a
was
no i o c l
( ,
-:
o./o;
//
- zl; /)
= o.ooo2;
l:ig c
).
So c cha qcs
c c
o l-ccs
l i
hc l u
hi l
[) ,c iously
oos cd
in, and
o l c s wc c
() ccs
hi c
hacl
o
ackccl
l
i cli iclual
o bc o c.
Mo lal
nu n c
o{'dil'lc cn
ccs
sc l
l
cacl
l a
also
was
only wo,
u
incli i i als
sc l
i onc o li u .
N l>c
o -clil'lc c
ccs
uscd
by cac6
[ a
als. was
co cli cd
wi l ' c
il'obsc a i.
iays
( , = o.z ;
l - 2.1;
/.
= l.<l lc ;
I i c
).
A c agc
dis ance
bl all
j7 oos
changcs
was
24.2
1.6
(l_ 7B)
n .
A nong
a lul s,
clis ancc
o ccl
as
inclcPc l
ol- cp ocl c i c
concli ion
(X = o.78; l.l'.
- p = <' .( 7),
and hc c
l i s
no dilli' cnce
bc wccn
aclul
lc alcs
a <l
ju e ilcs
(z = o.z9; p = o.l7);
jis a6cc
ng cd
as
inclepe ,
lcn o -yca
(X = .1.o,1;
l. . = ,- p = o.zz). Wc c
[a s si 'ply . o lns o
l c
lca cs
i ail.l[ lc
cc?
I1'so,
l c
lis .ib ion
o 'oos
changc
dis anccs
sli lL lci
[lc hc sa c
as hc clis iLlu ion
< l
(lis .u ccs
[ c wccn., ch
oos
ancl
i s
l ei cs
cighboL
in ha
yca
([]igL c
z).'l'he o,
l owc e ,
di -
c ecl
signi ican ly
(Fisl e 's
Exac
li:s :
p = (.).ool).
Al hough
67(yo
o
" i hi -yca ,
nei cs - ciglibclu
dis anccs
wc c
(5o n,
< nly
7 il
o oos
change
clis a nces
e c
ha
.small.
we co npa cd
he
e . clency
o changc
oos s
iln ong
adul
cn ales
using
a
-l
x 2 con ingcnc
ablc;
l c
h cc
colu ns
c[) cscn ccl
cp oduc i c
condi ion,
and he ows
ep esen ed
he numbe
o[ occasions
ha
a ba
was
obse ecl
in he
sa ne
oos
as
on he p e ious
day e sus
a oos
di -
c en
o n
ha uscd
hc p c ious
obse a ion
day.
Roos -swi ching
as
no andon l
clis ibL cd
an ong
cp ocluc i c
condi ions
(X = 6. iz;
d. .= ; P <o.ol). Fo
p egnan
ba s,
we eco ded
9 days
on which
a
oos
changc
occu cd
and zo ciays
on which i di l no , indica ing
. 49(il
chance
ha
a p egnan
ba would no bc in he
samc
oos
on consecu e
days.
Fo
lac a ing
and pos -lac a ing/non- ep oduc i e
en ales,
we
eco ded
7 changes
e sus
zB
non-changes (zoo/o),
and 6 changes
e sus
13
non-changes (3zo/o),
espec i ely.
Ju eniles
swi ched
oos s
on 5
days
and
emaincd
i' he samc
ee
on o days
( l%), a equency
i ually
iden i-
cal
o ha
o he pos -lac a ing
emales
(X2 = o.o ;
d. .= ,; p >o.i).
'[b in es iga e
whe he
oos -swi ching
was
ela ed
o ambien
empe a_
u e, we
cxamincd
he
co ela ion
be ween
oos -cmc gence
coun s
and
minimum a bien
empe a u e
he
day
o lie
obsc a ion
(usually
.ccu -
ing nca
clau'n hcn
l a s
a e
selec ing
il o()s )
. n i
n . x - ]un)
cn pc a-
u c
on hc clay
bc o e
hc
co , .
( hich
migh
cle c ninc
he he
ba s
e u n he
nex
o ning).
' : npe a u cs
we c
ob aincd
o n
a eco ding
s a ion
a
challo e,
l4 kn ESE
o ' he
s udy
si c.
,Ve
es ic ed
ou
analy-
sls o ees
a hich
we n aclc
) o eme gencc
coun s
in any
one
yea ,
a e
14
i 4ay,
bu belo e
17
July,
which
as
he
ca lies
cla e
ha
olan
ju eniles
,e c
cal) l cd.
Vc
Pc lo 'n cd
i e
analyses
on ou oos
ecs,
bu he e
was
no consis cn
ela ionship
l e wccn
he
en i onmcn al
a iables
and
loos
popula ion
sizc.
' o
ccs
sho ed
signi ican
co ela ions
bc ween
popula ion
sizc
i cl
bo h c nPc a u cs,
wo oos s
hacj
no signi ican
co -
.cla ions,
a lcl
l lc
clispllyccl
a signi ican
cla i, ship
i I linimun
e n,
pc a u c,
[ u
no naxi . .. - n
('lablc
I
).
sECT oN
l?oc s/i c loo
Roos -changing
Beha iou
=
0./0
P
=
0.0002
n
=
23
ba s
a
C]
c
c
U
=
0.71
p
<
0.0001
n
=
2l ba s
o
@
F
4567
Obse a ion
Days
Roos -chonging
beho iou .
(Top) The elo ionship
be ween numbe o
oos
chonges
ond numbe o doys ha he bo was loco ed. (Bo om)
The
elo ionship
be ween numbe o di e en ees used ond numbe o
doys ho he bo wos loco ed.
o7
-
o
l
o
i
Roos -change
e sus Nea es -neighbou
Dis ances
20.0
17 .5
5.0
12.5
0.0
7.5
5.0
2.5
0.0 101-1s0
Dinance
(m)
F cuRE z Roos -chonge
dis onces compo ed
o nea es -neighbou
dis ances wi hin
ho yeo .
Neo es -neighbou
dis ances
we e
meosu ed
only
o ees o
which
lndiono
bo s
we e
oc uolly odio- ocked in ha yeo .
We made eme gence
coun s on 16 da es,
obse ing l-Z oos s
on any
gi en
nigh , o a o al o 283 obse a ions.
On u3 occasions
(4oolo)
no
ba s
le he oos ;
occasionally he lack
o ba s
was
due o cold o we
wea he ,
bu we a ibu ed
mos ins ances
o he ba s' endency
o concen-
a e
ac i i y a some
ees and a ely
occupy o he s
(see
below).
Du ing
zo o he
emaining
7o nigh s
( zo/o),
only a single
ba eme ged.
Twel e
obsq a ions o soli a y e mc gence we e
o se en ba s ca ying ans-
1'A -E Ronk
co elo ions
be ween
numbe
o bo s lea ng
o ee ond
he
moximum
ombien empe o u e eco ded
on he p e ious
doy ond he minimum
ambien empe o u e
eco ded
on he
mo ning
o he coun .
Popula lon Slze and
Rela l e lmpo ance
o Roos s
l{oos /yea n Rangc
o da es Maximum
en pe a u e
(oC) Minimum
e npc a u e
("C)
24, 1992 4
zEl 992 l3
2Al 993 25
3Bl 1993 l(
3F 1993 27
30 May-15
July I
16
Junc-
l0 )uly
19 May-10
July
I |une-
15
)uly
14
unc-I4 uly
= -0.27;
p = 0.34
= 0.50;
p = 0.08 ,
= -0.
l{ ;
p = 0.38 ,
= 0.70;
p = 0.003 ,
= -0.56;
p = 0.007
0.02.;
p = Q.95
0.56;
P = 0.047
-0.
15;
p=Q.43
0.84;
P=0.000
I
-0.59;
p = 0.004
Fis e 's
Exac
Tes
p-0007
I cnung"
l Nea es
sECT oN
Roos ng lo8
mi e s,
so we
k.ew ha
a leas
one
ju enile
and h ee
p egnan ,
one lac-
a ing,
and
wo pos -lac a ing
{'e nales
oos ed
alone
on a leas
one nigh .
The o he
l5o
eme gc ces
in ol ecl
z-45
ba s, and mos
(g9%o)
in ol ed
z-z ba s
(Figu e
3).
- io
cie e nine
whc he
oos
popula ions
e e
consis en ly
la ge
a e
young
became
olan , c
con pa cd
eme gencc
coun s
made
be o e
and
a e 7 July,
in 99z
and 1993,
he
wo yea s
du ing which ield
wo k
sp. nnccl
hc
cn i c
sp ing-sun nc
oos ing
pc iod
(Figu c
4).
To
mini_
nize
co nplica ions
esul ing
om
mig a ion,
we
also
limi ed
he
da a
o
coun s
n. ade
be wee
14
May and 15 Augus .
The e
was
no sieni ican
di _
e encc
in popula ion
size
be o e
anci
a e
y July,
in ei hc
99z
lz - o.92; p = c36) o 99l (z = .o6;
p = o.z9).
This
sugges ed
ha Indi-
ana
ba s
occupied
a g ea e
numbe
o ees wi hin he
we land
soon
a e
he
young
became
olan ,
o ha
so ne
dispe sed
o o he
a eas.
Based
on eme gence
coun s,
we
es ima ed" he
ela i e
impo ance
o
oos
ees
by calcula ing
he
numbe
o "ba -days"
ha
each
was
occu-
pied;
o'e ba -day
equals
one
ba using
a ee
o one
day.
Indiana
ba s
did no use
each ee
o he
same
ex en ,
bu seemed
o p e e
wo o ou
pa icula
ees
e e y yea
(Figu e
5).
In 1993,
o exa ple,
ou oos s
( o, 2,a.,
3n,
and
3 ), ou o 8 ha
we e
occupied,
accoun ed
o 579
(8o%) o Tz ba -days;
in con as ,
eigh
o he
ees
( n, e, zF,3D>
3 ,3L
3 -,
and
3 )con ibu ed
a o al
o
only
4 eo/o)
ba -days.
Du ing
he
Eme gence
Coun s
(199'l
-93)
22-31 32-41 42-51
C oup
Size
; (;L ,
3 His og om
showinq
' equency
o oos
populo ion
sizes
CAN
,
o
a
109
TITERATURE
CITED
Anonyn ous.
995.
A c
An c ican
L a s losing
g o ncl?
Ba s
3:,1.
Ba bou ,
Il.W.
a'd W.ll. Da is.
9( 9.
Bi s
.{' A'c ica. Uni e si y
oI
Kcn ucky
l) css,
Lcxing on,
Ky.
B igham,
R.M.
99 . Flcxibili y
i. b agi.g
a'd oos i g
bcha iou
by hc
big
b own l
(L:.p csicus
usc s
). (.an.
l. Zool. 6g: 7-21.
Calahan,
I . ., III. 99j.
Indiana
l ii su n ne
l habi a
cqui emcn s.
M.s.
'l'hesis,
Uni c si y
o Missou i,
Columbia,
Mo.
clawson,
I . 9i 7.
I.cliana
ba s:
clown lb hc coun .
I]ndange ed
species
Tcchnical
Bulle in
22..9-11.
Fos e ,
R. 991.
Roos
si c sclcc ion
by he no hc n
ba ,
A4yo is
sep_
cn ionulis.
Ba
Res.
News
-14: o8.
Ga dnc ,,.8.'
J.D.
Ga ne , and
J.E.
Ilo inann.
99 .
Sum ne
oos
selec-
ion and
oos ing
bcha io
o Myo is
sodalis
(lndiana
ba ) in
IIIin.is.
U'publ. cp.
Illinois
Na u . l
His o
Su cy,
champaign,
Il
IJa '
J.s.
97 .
Roden s.
In con pa a i c
ph siclogy
o he mo egula ion,
Vol.
z.
G.C. Whi ow
(cdi o ).
Academic
P ess,
No yo k, N.y.,
PP.
- 49.
He nshaw,
R.E,. and
G.E. Folk,
J .
y66.
Rela ion
o[ he mo egula ion
o
seasonally
changing
nic'oclima e
in o
species
o ba s
(Myo is
Iuci ugus
and M. sodalis).
physiol.
Zool. 39:24-36.
Humph ey,
S.R.,
A.R.
Rich e ,
and j.B. Cope.
y77.
Summe habi a and
ecology
o[ l. e
enda'ge ed
Indiana
ba , Myo is
sodalis.
I. Mammal.
58334-36.
Kunz,
T.H.
1982.
Roos i'g
ecology
o ba s.
In Ecology
o[ ba s.
T.H. Kunz
(edi o ).
Plenum
P ess,
New
yo k,
N.y.,
pp. _j .
Ku a,
A. 995.
Ba k
oos
oIa male
big b own ba (Ep esicus
uscus).
Ba
Res. News
35:63.
Ku a,
A. and R.
Fos e .
1995.
The
b own
c ecpe
(A es:
Ce hii<1ae):
a
compe i o
o ba k- oos ing
ba s?
Ba
Rcs.
News
36:6_7.
xu a, A. and
T.H.
Kunz.
1987.
Size
o ba s
a bi h and ma e nal
in es -
nen
in p egnancy.
Symp.
Zool.
Soc.
i.ondon 57:79_ o7.
London,
U.K.
Ku a,
A. and
K.J. Williams.
994.
Thc mal
aspcc s
o Indiana
ba s oos _
ing
in ees.
Ba
Res.
News
35: o4.
Ku a,
A., D. King,
J.A.
Te amino,
I.M. S iblcy,
and K.J. Williams.
993a.
Summe
oos s
o he
endange .ed
Indiana
ba (Myo is sodalis) on
he no he n
edge
o i s ange.
A ne .
Midl. Na .
iz9: 3z_3g.
Ku a,
A.,
I. Ka h,
E.L.
S ni h,
Il. Fos c ,
N4.
O ick,
and R. Ross. 993b.
A
ma e ni y
oos
o he
e'dange ed
I'diana ba (Myo is sodalis)
in a
unshaded,
hollow,
sycamo e
ee
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occiden ali.s).
Ame .
Midl.
Na .
3o:4o5-o7.
s ECTI
o N Roos ing
Lewis,
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ideli y
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I. Mammal.
76:4g *96.
Lomolino,
M.V. and R.
Channeli.
995.
Splendid
isola ion:
pa e ns
o
geog aphic
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collapse
in endange ed
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J.
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76335-47.
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A.G.
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Ecology
o inscc s
pa asi ic
on ba s. In Ecology
o
ba s. T.H.
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J.
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S.R.
Humph ey,
J.B.
Cope,
and
V. B ack,
J .
1993.
Modi ied
ca e en ances:
he mal
e ec
on body mass
and esul ing
decline
o
endange ed
Indiana
ba s
(Myo is sodalis).
Conse a ion
Biology
7i4o7- 5.
S udie , E.H. 98 . Ene ge ic
ad an ages
o sligh
d ops
in body empe a-
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