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In aspeci ic Va ia ion in Roos -si e Selec ion
by Li le B own Ba s (Myo is luci ugus)
. .
ABSTRACT
Al hough many species o ba s selec oos si es in la ge ees ha a e in
open a eas, in aspeci ic a ia ion in oos -si e selec ion may exis . We col-
lec ed da a on he oos ing beha iou o li le b own ba s in he Cyp ess
Hills, Saska chewan, o de e mine he ex en o in aspeci ic a ia ion in
oos -si e selec ion. In addi ion, we examined he he mal mic oclima e o
he ee- oos s selec ed by ba s, o de e mine i oos -mic osi e a ia ion
can explain why ce ain ca i ies a e selec ed o e o he s. We ound li le
b own ba s oos ing in ees as well as buildings. Wi h he excep ion o a
male who oos ed in a sp uce (Picea glauca) s ump, ee- oos s selec ed
by male and emale li le b own ba s we e all in embling aspen (Populus
emuloides) ees. We ound a ia ion in oos -si e ideli y and di e en ial
use o o po by male ba s. Tempe a u es wi hin coni e snag ca i ies di -
e ed om aspen ca i ies du ing he day, and mi o ed ambien empe a-
u e, which ended o be wa me han aspen ca i ies. We p opose ha
ba s selec ca i ies in aspens because hey a e suscep ible o hea o .
Aspen ees wi h hea o p o ide ca i ies wi h an in ac sapwood shell
ha p o ec s ba s agains ha sh ambien condi ions as well as p eda o s,
and p o ides a unique he mal mic oclima e. Ou esul s sugges ha he
o igin o a oos si e may be unimpo an o a ba , p o ided ce ain o he
equi emen s a e me .
INTRODUCTION
Compa ed wi h nes -si e selec ion by bi ds, ela i ely li le is known abou
oos -si e selec ion by ba s, especially selec ion o na u al oos s, such as
ee ca i ies. Ra he , he majo i y o ecological esea ch on empe a e
insec i o ous ba s (Chi op e a: Vespe ilionidae) has deal wi h species
oos ing in human-made s uc u es. The eason o his is ha i is a
easie o ind and gain access o ba s oos ing in a human-made s uc u e
han in na u al si es, such as ee ca i ies. As a esul , howe e , ou unde -
s anding o he cues ha ba s use o selec si es unde na u al condi ions,
whe e human-made s uc u es a e no abundan , is limi ed.
Un il e y ecen ly, he numbe o s udies ha examine selec ion o
***Rep in ed wi h pe mission. No u he ep oduc ion is au ho ized wi hou w i en pe mission om C own Publishing G oup.
This e sion o he documen is no he e sion o eco d. Figu es and/o pic u es may be missing om his o ma o he documen .***
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Roos ing
ee- oos si es by empe a e ba s has been low. Howe e , as e idenced
by he numbe o pape s p esen ed a his con e ence ha add ess ee-
oos ing by ba s, his is clea ly changing (see pape s in his olume by
Be s, Chung-MacCoub ey, C amp on and Ba clay, Ku a e al., O msbee,
Sasse and Pekins, and Vonho ). O e di e en o es ypes and geog aphi-
cal egions, in o ma ion on he cues ha ba s use o selec oos si es
unde na u al condi ions is apidly eme ging. Despi e di e ences in s udy
a eas and species, oos ees a e ypically la ge and in open a eas. In
addi ion, many ba s ha oos in aspen ees a e seconda ily using ca i ies
o iginally exca a ed by p ima y ca i y exca a o s (sapsucke s and o he
woodpecke s) o nes si es.
Al hough we hea d a he con e ence ha many species o ba s selec
oos si es in la ge ees ha a e in open a eas, in aspeci ic a ia ion in
oos -si e selec ion has no been a ocus o a en ion. Howe e , in he
Ne he lands, he noc ule ba (Nyc alus noc ula) is ound exclusi ely in
ee ca i ies, while in cen al Eu ope indi iduals oos in buildings as well
as ee ca i ies ( an Hee d and Slui e ). Big b own ba s in B i ish
Columbia oos in hollows o dead ponde osa pine ees (Pinus ponde -
osa), while he same species in On a io oos s in human-made s uc u es
(B igham ). Indeed, geog aphic di e ences, such as clima ic condi-
ions and p ey a ailabili y, may explain di e ences seen in he ype o
oos selec ed. This leads o he ques ion o how lexible he oos ing
beha iou o ba s is in one a ea, whe e geog aphic di e ences a e no a
ac o .
We collec ed da a on he oos ing beha iou o li le b own ba s in he
Cyp ess Hills, Saska chewan, o de e mine he ex en o in aspeci ic a ia-
ion in oos -si e selec ion. In addi ion, we examined he he mal mic o-
clima e o he ee- oos s selec ed by ba s o de e mine i oos -mic osi e
a ia ion can explain why ce ain ca i ies a e selec ed ela i e o o he s.
We p opose ha ba s selec ca i ies in aspen ees in ec ed wi h hea o
because hese ees p o ide ca i ies wi h an in ac sapwood shell ha p o-
ec s agains ha sh ambien condi ions, as well as p eda o s, and p o ides
a unique he mal mic oclima e wi hin.
METHODS
Ou s udy occu ed om May o Augus , . The s udy a ea was loca ed
wi hin he Wes Block o Cyp ess Hills P o incial Pa k (° ′
, °
′
), app oxima ely km sou hwes o Maple C eek, Saska chewan,
Canada. Lodgepole pine (Pinus con o a) o es , wi h li le unde s o ey,
occupies d y si es abo e m, and whi e sp uce (Picea glauca) o es
occu s in cool, mois a eas nea we lands and on no h- acing slopes.
T embling aspen is ound g owing wi h whi e sp uce nea s eams as well
as in s ands sca e ed h oughou he hills on he edge o lodgepole pine
s ands (Sauchyn ).
Va ia ions in
Roos ing Beha iou
by Li le B own Ba s
Da a on oos -si e selec ion in buildings by li le b own ba s we e col-
lec ed by wa ching he buildings in he s udy a ea a dusk, wi h he aid o
a ba de ec o , o de e mine i ba s we e oos ing in he building. Da a on
oos -si e selec ion in ees by li le b own ba s we e collec ed as pa o a
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la ge s udy looking a na u al oos -si e selec ion by big b own ba s
(Kalcounis ). Li le b own ba ee- oos si es we e ound using wo
me hods. Fi s , while aking measu emen s o ca i y mic oclima e o big
b own ba (Ep esicus uscus) oos and andom a ailable ees (Kalcounis
), we checked ca i ies o li le b own ba s. Second, we used adio-
eleme y o loca e oos ees. Indi idual li le b own ba s we e cap u ed
in mis ne s se in suspec ed o aging a eas. Upon cap u e, .-g, empe a-
u e-sensi i e ansmi e s (
-, Holohil Sys ems L d., Ca p, On a io)
we e a ached using Skinbond cemen . Ba s we e eleased soon a e he
cemen had d ied. Indi iduals ca ying adio- ansmi e s we e acked o
hei oos s on he mo ning ollowing cap u e using Me lin (Cus om
Elec onics, U bana, Illinois) po able eleme y ecei e s and hand-held,
-elemen Yagi an ennae. When a suspec ed oos ee was loca ed, we
obse ed i a dusk o ensu e ha i was being used by ba s, and o de e -
mine he numbe o ba s oos ing in he ee. Ba s ca ying adio- ans-
mi e s we e acked o hei oos s e e y day un il he ansmi e ell o
o he signal was no longe hea d. Fo each day ha we acked ba s, we
de e mined i hey we e ac i e o o pid in he oos .
The mal Mic oclima e
Wi hin Ca i ies
To compa e empe a u e p o iles o he ca i ies selec ed by ba s (old sap-
sucke holes in aspen ees) wi h hose no selec ed (ca i ies in coni e
snags), empe a u e-sensi i e adio- ansmi e s we e used. Snags we e
de ined as dead whi e sp uce o lodgepole pine ees wi h ca i ies. T ans-
mi e s we e placed in ca i ies by climbing he ees and suspending he
ansmi e in he ca i y using mono ilamen line. Two ca i ies o each
ype we e selec ed a andom.
Tempe a u es in bo h ca i y ypes we e compa ed o ambien empe a-
u es. Each sampling day was di ided in o ou ime pe iods e lec ing
ea ly mo ning (pe iod : h–h), mid-mo ning (pe iod : h–
h), a e noon (pe iod : h–h), and e ening (pe iod : h–
h) ime blocks.
To measu e empe a u es, we used an au oma ed eleme y ecei e
(Lo ek
eleme y ecei e using E en Log Ve sion .,
da a-
logging so wa e, Lo ek Enginee ing Inc., Au o a, On a io). The ecei e
was p og ammed o eco d he empe a u e, o each ansmi e , e e y
hou o each -hou pe iod om o July. To de e mine whe he
ambien and ca i y empe a u e p o iles di e ed, we pe o med one-way
analysis o a iance (
) es s using mean empe a u es wi hin he
ou ime pe iods.
RESULTS
Roos -Si e Selec ion
by Li le B own Ba s
We ound wo li le b own ba building- oos s, bo h o which we e ma e -
ni y colonies. One ma e ni y colony was loca ed in he pa k headqua e s
building in he Wes Block o Cyp ess Hills P o incial Pa k. The ba s
oos ed in he a ic and unde he ceda shingles o an eas - acing sec ion
o he oo . They ha e used he s uc u e o a leas ou yea s (see
Kalcounis and B igham ). The mean colony size in and was
wi h a ange o o (Kalcounis and B igham ). The o he
building- oos was loca ed in a a mhouse abandoned o one yea . The
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Roos ing
a mhouse had b oken windows h ough which he ba s eme ged. Du ing
an eme gence in June , o e li le b own ba s we e coun ed.
We ound six li le b own ba ee- oos s, one o which housed a
ma e ni y colony. While aking measu emen s wi hin he ca i y o a an-
dom a ailable ee o Kalcounis (), a colony o li le b own ba s
was ound in an old sapsucke (Red-naped o Yellow-bellied subspecies o
Sphy apicus a ius) hole in a li e embling aspen ee. The wo indi id-
uals cap u ed om his colony we e ju enile emales oo small o ca y
ansmi e s. The colony emained in he oos ee o h ee days.
The emaining i e oos si es we e loca ed by adio- acking wo adul
male li le b own ba s. The wo ba s we e caugh on and July, espec-
i ely, which allowed us o adio- ack hem simul aneously. Bo h males
oos ed soli a ily. We we e able o ack he i s male ba o eigh con-
secu i e days, and ound ha i oos ed exclusi ely in a c ack in a sp uce
s ump. We we e able o ack he second male ba o consecu i e days
and ound ha i oos ed exclusi ely in aspen ees; howe e , i swi ched
be ween ou di e en ca i ies in ou di e en ees. De ails o he male
li le b own ba oos - ee cha ac e is ics a e gi en in Table . All aspen
ees had symp oms o ungal hea o in he o m o conks ( ungal
ui ing bodies).
In addi ion o a ia ion be ween he males in he ype o oos selec ed
and ideli y o pa icula oos si es, he males di e ed wi h espec o he
use o o po while in he oos . O he eigh days ha we we e able o
ack he males simul aneously, bo h ba s emained ac i e in he oos o
se en days. On one day, howe e , he s ump- oos ing male emained ac i e
while he o he male oos ing in a li e aspen ee used o po (Figu e ).
The mal Mic oclima e
Wi hin Ca i ies
Mean empe a u es di e ed signi ican ly in ime pe iods (F = .,
d = , p <.), (F = ., d = , p <.), and (F = .,
d = , p <.). Roos ca i ies we e signi ican ly wa me han ambien
empe a u e, bu signi ican ly coole han coni e snags du ing ime
pe iod (Figu e ). In ime pe iod , oos ca i ies we e signi ican ly
Cha ac e is ics o male li le b own ba ee- oos s. The male ba who oos ed in he sp uce s ump is e e ed
o as Ba A. The male who swi ched aspen ee- oos s is e e ed o as Ba B.
Ca i y
T ee en ance Numbe
T ee T ee heigh O igin heigh o ca i ies Da es
Ba condi ion species (m) (cm) o ca i y (m) in ee occupied
A dead, s ump sp uce 2.2 38.2 spli wood — — 6–14 July
B dead, snag aspen 6.1 33.3 sapsucke 5.7 3 8 July,
14 July
B li e aspen 31.2 35.0 sapsucke 12.0 1 9–10 July,
16 July
B li e aspen 39.4 36.7 b anch sca 7.7 1 11–13 July,
15 July
B dead, s anding aspen 12.6 25.1 sapsucke 8.1 >5 17–22 July,
27 July
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40
33
26
19
12
5
Tempe a u e (°C)
1200 240 360 480 600
Time (min; 0=1300 h . CST)
Tempe a u e p o iles o he wo male ba s. The ba oos ing in he
s ump (•) s ayed ac i e while in he oos . The ba oos ing in he
aspen ca i y (
O
) was o pid un il app oxima ely 1515 h and was ac i e
by 1900 h. Ambien empe a u e is shown by he solid line.
27
25
22
20
17
15 1234
Time Pe iod
Mean Ca i y Tempe a u e (°C)
Mean (
±
1 SE) empe a u es o ca i ies in each o he ou ime pe iods
(see ex ). Means sepa a ed by * a e signi ican ly di e en (Tukey’s
Tes ). Ambien (
O
), Roos (•), Snag ( ).
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Roos ing
wa me han ambien empe a u e and snags, which did no di e om
one ano he (Figu e ). In ime pe iod , bo h ca i y ypes we e signi i-
can ly wa me han ambien empe a u e (Figu e ).
DISCUSSION
Ou esul s demons a e ha wi hin an a ea he e is conside able in a-
speci ic a ia ion in li le b own ba oos -si e selec ion, wi h ba s oos ing
in bo h li e and dead ees as well as buildings. Wi h he excep ion o he
sp uce s ump, he e was li le a ia ion in he ype o ee- oos selec ed
by male and emale li le b own ba s. We ound a ia ion in oos -si e
ideli y and di e en ial use o o po by male ba s. These esul s sugges
ha he o igin o a oos si e may be unimpo an o a ba , p o ided ha
ce ain o he equi emen s a e me . Indeed, he oos ing beha iou o li -
le b own ba s in Cyp ess Hills is lexible enough ha hey we e ound
oos ing in buildings, as well as aspen ees. P esumably, li le b own ba s
a e able o use human-made s uc u es o oos s, p o ided ha he
human-made s uc u es a e o app op ia e dimensions and p o ide a sui -
able mic oclima e wi hin.
Ba s used mo e han one oos ee. The e a e se e al easons why
some o he ba s in ou s udy may ha e swi ched ee- oos s. When he
oos mic oclima e o dis ance o o aging a ea changes, ba s may espond
by swi ching o mo e p e e ed si es (Kunz ; Lewis ). Ba s may
swi ch oos s o a oid p eda o s (Ba clay e al. ; Kunz ; Wilkinson
; Aude ; Lewis ; bu see Fen on and Rau enbach ) o
la ge popula ions o ec opa asi es (Wilkinson ; Lewis ).
Some species o ba ha swi ch oos s equen ly emain ai h ul o
pa icula a eas (Lewis ). E en hough one male li le b own ba
swi ched oos s equen ly, he o en e u ned o p e iously used oos
ees, sugges ing ideli y o a pa icula g oup o oos ees a he han a
single oos ee. Big b own ba s in he Cyp ess Hills exhibi simila oos -
swi ching beha iou (Kalcounis ), as did ba s in many s udies p e-
sen ed a he con e ence (see pape s in his olume by Be s, C amp on
and Ba clay, Ku a e al., Vonho ).
Wi h he excep ion o he sp uce s ump, all ee- oos s we e in aspen
ees. Mos ca i ies wi hin he aspen had been exca a ed and used as nes
si es by sapsucke s. Sapsucke s and o he woodpecke s a e p ima y ca i y
exca a o s. The use o aspen by p ima y ca i y exca a o s is ela ed o he
ela i ely so wood and suscep ibili y o hea o o hese ees. False in-
de o (Fomes ignia ius) is a majo cause o decay in aspen (Basham
; Pe e son and Pe e son ). False inde o in ec s aspen h ough
oo s o b oken b anches (Basham ), and induces ex ensi e decay o
he hea wood while spa ing he sapwood, which emains as a ough, li -
ing, ou e shell (Kilham ). Hoo -shaped conks a e cha ac e is ic ex e -
nal indica o s o alse inde o (Pe e son and Pe e son ), and may be
he cues ha p ima y ca i y exca a o s use o selec aspen wi h decayed
hea wood. Thus, in he absence o sapwood o , an aspen in ec ed wi h
alse inde o p o ides ideal condi ions o nes ing p ima y ca i y exca-
a o s ha a e capable o ge ing h ough he sapwood laye . In ec ed
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aspen ees p o ide ca i ies wi h a shell ha p o ec s agains ha sh, ambien
condi ions, as well as om p eda o s no s ong enough o chew h ough
he sapwood laye o small enough o i h ough he en ance hole.
Ca i y- oos ing ba s, like a ious bi ds, a e seconda y ca i y use s. As
seconda y ca i y use s, ba s choose ca i ies om among hose al eady exca-
a ed, and a e he e o e cons ained by he p ima y ca i y exca a o s’ p e -
e ence o nes ing si es and he decay cha ac e is ics o he ee and ca i y.
The selec ion o nes si es by p ima y ca i y exca a o s p obably in luences
he popula ion numbe s and communi y composi ion o ba s in he
Cyp ess Hills, jus as he local dis ibu ions and abundances o se e al sec-
onda y ca i y-nes ing bi ds a e enhanced by old nes ca i ies (Daily ).
P e e ence o aspen ees has been shown o many p ima y ca i y-
exca a ing species. Th oughou he No h Ame ican b eeding anges o
sapsucke s and o he woodpecke s, mos o he exca a ed nes holes a e in
aspen ees (A izona, Li and Ma in ; Colo ado, C ocke and Hadow
, Win e ni z and Cahn ; New Hampshi e, Kilham ; B i ish
Columbia, E ksine and McCla en , Pe e son and Gau hie ,
Keiske , Ha es ad and Keiske ). E en when aspen is no he
dominan ee species in a o es , p ima y ca i y exca a o s choose i o e
o he species wi h a highe a ailabili y. In he In e io Douglas- i Biogeo-
clima ic zone o sou he n B i ish Colombia, .% o all p ima y ca i y-
exca a ing bi ds we e ound nes ing in aspen ees, despi e a ela i e a ail-
abili y in a andom sample o only .% (Ha es ad and Keiske ). As
pa o a s udy ha examined ep oduc i e success o ca i y-nes ing bi ds,
aspen ees p o ided g ea e han % o all nes si es o p ima y ca i y
exca a o s, e en hough aspen cons i u ed only % o all ee species in
andom plo s (Li and Ma in ).
In he Cyp ess Hills, sui abili y and selec ion o aspen o nes ing, and
subsequen oos ing, seem o be de e mined bo h by hei a ailabili y as
he only dominan ha dwood, and by decay cha ac e is ics. Selec ion by
ba s o aspen ees o e whi e sp uce and lodgepole pine is no su p ising
gi en he di e ence in decay cha ac e is ics o he ees. Whi e sp uce and
lodgepole pine a e so woods, which do no ha e he same decay cha ac-
e is ics o ha dwoods, such as aspen. In coni e s, hea wood and sapwood
bo h decay mo e apidly, which p ecludes he o ma ion o a solid ou e
shell o sapwood (McClelland ).
P ima y ca i y exca a o s in he Cyp ess Hills a e limi ed in he numbe
o species o ee ha hey can selec o exca a ion, as aspen is he only
abundan ee wi h sui able decay cha ac e is ics. The decay dynamics o
balsam popla s (Populus balsami e a) a e simila o aspen; howe e , balsam
popla s a e no abundan in he Cyp ess Hills. Whe e he dis ibu ion o
aspen o e laps wi h ha o o he ees wi h simila decay cha ac e is ics,
such as wes e n la ch (La ix occiden alis) and pape bi ch (Be ula papy i-
e a) in he Rocky Moun ain o es s o no h-wes e n Mon ana, p ima y
ca i y exca a o s p e e nes ing in wes e n la ch (McClelland e al. ).
Tempe a u es in aspen ca i ies ended o be wa me han ambien em-
pe a u e a nigh (pe iods and ) and coole han ambien empe a u e
du ing he day (pe iods and ). Du ing he day, empe a u es wi hin
coni e snag ca i ies di e ed om aspen ca i ies and mi o ed ambien
empe a u e, which ended o be app oxima ely °C wa me han aspen
ca i ies. The dicho omy in empe a u es be ween aspen ca i ies and
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Roos ing
coni e snag ca i ies sugges s ha i may be he e ec o wa m ambien
empe a u e ha in luences selec ion o aspen o e coni e snags.
The di e en decay dynamics o so woods and ha dwoods should in lu-
ence empe a u e egimes wi hin ca i ies o snags and aspen ees. Ca i ies
in coni e snags a e also a ailable as oos si es o ba s in he Cyp ess Hills.
Despi e hei a ailabili y, ca i ies in snags we e ne e used as oos si es.
The o ma ion o a solid ou e shell in aspen may no only p o ec ba s
om p eda o s, bu may also p o ide a bu e om high ambien empe a-
u es. Decaying coni e s do no o m his solid ou e shell.
Bu ne and Augus ’s () s udy on he ene gy budge s o a ma e ni y
colony o building- oos ing li le b own ba s o e s insigh as o why he
day ime di e ence be ween aspen and snag ca i ies may be an impo an
c i e ion in he selec ion o oos si es by li le b own ba s. A noon,
unoccupied li le b own ba oos s in hei s udy we e °C as compa ed
wi h °C o occupied oos s. The he moneu al zone o li le b own
ba s is be ween .°C o .°C, which pu s he empe a u e o occupied
oos s wi hin he he moneu al zone. By occupying a oos , li le b owns
inc ease he empe a u e by °C. I ollows hen, ha an unoccupied oos
ha is much wa me han °C would be unsui able o li le b own ba s
because once occupied, he empe a u e wi hin he oos would app oach,
o exceed, he uppe le el o he he moneu al zone. Fo his eason, we
sugges ha he empe a u e di e ence be ween aspen and snag ca i ies
du ing ime pe iod ende s snag ca i ies less sui able as li le b own ba
oos si es. To es his, empe a u e p o iles o aspen ca i ies occupied by
li le b own ba s a e equi ed.
ACKNOWLEDGEMENTS
We hank Robe Ba clay and Ma k B igham o he in i a ion o p esen
ou wo k. Da id Gumme assis ed us in he ield, and we a e especially
g a e ul o him o leading us o he oos ee ha we ‘‘misplaced.’’ We
alue he discussions ha one, o bo h, o us ha e had wi h Da en
Bende , Paul B adshaw, Lisa C amp on, Sco G indal, Da id Gumme ,
and Maa en Vonho ega ding his p ojec . We app ecia e he use o he
Depa men o Biology, Uni e si y o Regina Field S a ion, and acknowl-
edge he hospi ali y o he Wes Block o Cyp ess Hills and Fo Walsh
pa k adminis a o s and employees. Financial suppo was p o ided by
g an s om he Regina Na u al His o y Socie y, he Theodo e Roose el
Memo ial Fund (Ame ican Museum o Na u al His o y), and he Na u al
Science and Enginee ing Resea ch Council o Canada (
) in he
o m o esea ch g an s and Uni e si y o Regina P esiden ’s
g an s
o Ma k B igham.
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