Quan i a i e compa ison o ee oos s used by
ed ba s (Lasiu us bo ealis) and Seminole ba s
(L. seminolus)
Michael A. Menzel, Timo hy C. Ca e , B ian R. Chapman, and Joshua Lae m
Abs ac : We adio- acked 11 ed ba s (Lasiu us bo ealis) and 5 Seminole ba s (L.seminolus) o 64 and 34 day oos s,
espec i ely. Indi iduals o bo h species we e ound oos ing wi hin he canopy o he oos ees, clinging o lea pe ioles o
he ips o small b anches (<4 cm in diame e ). Red ba s oos ed p ima ily in ha dwoods (97%), whe eas he oos s o
Seminole ba s we e loca ed p ima ily in pines (94%). Ten o he 16 oos -si e a iables examined di e ed signi ican ly
be ween ed ba s and Seminole ba s: numbe o ees in he o e s o y, o e s o y heigh , unde s o y ichness and di e si y,
o e s o y ichness, di e si y, and e enness, oos - ee diame e , pe cen canopy closu e, and pe cen age o coni e s in he
o e s o y. These di e ences we e ela ed di ec ly o he di e en ial use o oos ing habi a s by he wo species. The oos s o
ed ba s we e loca ed in pine – mixed ha dwood communi ies and bo omland ha dwood swamps, while he oos s o
Seminole ba s we e loca ed in communi ies domina ed by pines. To examine wi hin-s and oos selec ion, he diame e ,
heigh , and species composi ion o oos ees used by ed and Seminole ba s we e compa ed wi h hose o neighbo ing ees.
Roos ees o ed and Seminole ba s had signi ican ly la ge diame e s and we e signi ican ly alle han su ounding ees.
Day oos s o ed and Seminole ba s we e loca ed in 18 and 5 ee species, espec i ely. The ee species used di e ed
signi ican ly om expec ed o he ed ba bu no o he Seminole ba .
Résumé : Nous a ons sui i pa adio élémé ie les déplacemen s de 11 Chau es-sou is ousses (Lasiu us bo ealis) e s 64
do oi s de jou e de 5 Chau es-sou is séminoles (L. seminolus) e s 34 do oi s. Les chau es-sou is des deux espèces on é é
ou ées dans le euillage des a b es des do oi s, c amponnées aux pé ioles des euilles ou aux ex émi és de pe i s ameaux
(diamè e <4 cm). Les Chau es-sou is ousses on é é ou ées su ou dans les bois du s (97%) e les Chau es-sou is
séminoles, su ou dans les pins (94%). Seize a iables eliées aux do oi s on é é examinées e dix d’en e elles di é aien
signi ica i emen chez les deux espèces : nomb e d’a b es dans l’é age supé ieu de la o ê , hau eu de l’é age supé ieu de la
o ê , ichesse e di e si é de l’é age in é ieu de la o ê , ichesse, di e si é e égula i é de l’é age supé ieu de la o ê ,
diamè e des a b es de epos, densi é du euillage en pou cen age, pou cen age des coni è es dans l’é age supé ieu de la o ê .
Les di é ences é aien di ec emen eliées à l’u ilisa ion di é en ielle des habi a s o e s pa les do oi s chez les deux
espèces. Les do oi s des Chau es-sou is ousses é aien si ués dans les communau és mix es bois du s – pins e dans les
ma écages bas à bois du s, alo s que les do oi s des Chau es-sou is séminoles é aien si ués dans les communau és dominées
pa les pins. Pou dé e mine les p é é ences de si es de epos à l’in é ieu d’un do oi , le diamè e, la hau eu e la
composi ion en espèces des a b es du do oi u ilisés pa le deux espèces de chau es-sou is on é é compa és aux mêmes
a iables des a b es a oisinan s. Les a b es u ilisés pa les deux espèces a aien un diamè e e une hau eu signi ica i emen
supé ieu s à ceux des a b es a oisinan s. Les do oi s des Chau es-sou is ousses compo aien 18 espèces d’a b es, ceux des
Chau es-sou is séminoles, 5 espèces. Chez les Chau es-sou is ousses, mais pas chez les Chau es-sou is séminoles, les
espèces d’a b es u ilisées di é aien signi ica i emen de celles indiquées pa les p édic ions héo iques.
[T adui pa la Rédac ion]
In oduc ion
Al hough ba s a e an impo an componen o o es ecosys-
ems, ou unde s anding o hei habi a equi emen s has
lagged behind ou unde s anding o hose o o he mammalian
g oups. Un il ecen ly, mos o he in o ma ion on ba oos s
and oos ing beha io came om obse a ions made a ca es,
mines, o a i icial s uc u es (Be s 1995). O he han anecdo-
al accoun s, li le was known o oos -si e selec ion among
ee- oos ing ba s (e.g., Ba clay and Cash 1985; Cons an ine
1958, 1966; Pa sons e al. 1986). Wi h he ad en o minia u -
ized adio ansmi e s, mo e de ailed analysis o chi op e an
habi a p e e ences, oos selec ion, and oos ing beha io is
possible (see Ba clay e al. 1988; Lunney e al. 1988; Taylo
and Sa a 1988; Vonho 1995). I is now easible o compa e
oos -si e selec ion be ween closely ela ed species in he same
locale (Lacki 1995).
In eas e n No h Ame ica, ew de ails a e known abou he
cha ac e is ics o ee oos s used by wo ela ed species, he
ed ba (Lasiu us bo ealis) and he Seminole ba (L. semi-
nolus). Based upon anecdo al accoun s, ed ba s a e assumed
o commonly oos in oliage a he edge o ha dwood ee
canopies (Ba bou and Da is 1969; Cons an ine 1966; Koon z
Can. J. Zool., 76: 630–634 (1998)
Recei ed July 3, 1997. Accep ed Oc obe 21, 1997.
M.A. Menzel, T.C. Ca e , and B.R. Chapman.1Daniel B.
Wa nell School o Fo es Resou ces, Uni e si y o Geo gia,
A hens, GA 30602, U.S.A.
J. Lae m.2Museum o Na u al His o y, Uni e si y o Geo gia,
A hens, GA 30602, U.S.A.
1Au ho o whom all co espondence should be add essed
(e-mail: Chapman@smokey. o es y.uga.edu).
2Deceased.
630
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and Da is 1991; McClu e 1942; Mum o d 1973). Thei oos s
may also be loca ed in clumps o Spanish moss (Tillandsia
usneoides; Cons an ine 1966; Jennings 1958), in coni e ous
ees (McClu e 1942), in woodpecke holes (Fassle 1975),
and unde sun lowe (Helian hus sp.) lea es (Downes 1964).
Seminole ba s o en oos in Spanish moss (Ba bou and Da is
1969; Cons an ine 1958; Ha pe 1927; Jennings 1958; Wilkins
1987). They may also occupy clumps o oliage (Sealande and
Heid 1990), ee b anches (Ba kalow and Adams 1955), he
ips o pine limbs (M.A. Menzel e al., in p epa a ion),3and
ca i ies unde loose ba k (Sealande 1979). Des uc ion o
oos s may be he mos impo an ac o in he decline o ba
popula ions in No h Ame ica (Kunz 1982). Because ed ba s
and Seminole ba s spend mo e han hal o each day in o es
oos s (Kunz 1982; Vonho and Ba clay 1996), conse a ion
o hese species may depend in pa on a de ailed knowledge
o hei oos -si e cha ac e is ics. Since hese wo species a e
closely ela ed, i is o en assumed ha hei oos loca ions a e
simila wi hin a limi ed geog aphic ange. Since s udies o
ela ed species in he same locale a e aluable o iden i ying
in e speci ic di e ences in oos -si e selec ion (Be s 1995),
we compa ed he cha ac e is ics o oos ees used by he ed
ba and Seminole ba in wo loca ions wi h simila o es cha -
ac e is ics.
Me hods
S udy a eas
We conduc ed his s udy du ing June and July 1995 and om June o
Augus 1996. The s udy was conduc ed on Sapelo Island, Geo gia, in
1995 and a he Sa annah Ri e Si e Na ional En i onmen al Re-
sea ch Pa k, Sou h Ca olina, in 1996. Sapelo Island is a 4411-ha
ba ie island loca ed app oxima ely 63 km sou h o Sa annah, Geo -
gia, and 5.5 km o he coas (31°27′N, 81°16′W). The Sa annah
Ri e Si e is a 76 900-ha Na ional En i onmen al Resea ch Pa k ad-
minis e ed by he Uni ed S a es Depa men o Ene gy. I is loca ed
23 km sou heas o Augus a, Geo gia (33°15′N, 81°40′W). Commu-
ni ies o longlea pine (Pinus palus is), loblolly and slash pine
(P. aeda and P. ellio ii, espec i ely) and pine – mixed ha dwoods
we e common o bo h si es. Pond pine (P. se o ina) communi ies and
ma i ime oak o es s domina ed by li e oaks (Que cus i giniana)
we e ound only on Sapelo Island. Bo omland communi ies we e
es ic ed o he Sa annah Ri e Si e.
Roos ing si es
Ba s we e cap u ed in mis ne s se up o e small ponds o c eeks o
along ails. The species, sex, mass, o ea m leng h, and age o all ba s
cap u ed we e eco ded. Indi iduals we e sepa a ed in o ju enile and
adul age-classes based on he deg ee o epiphyseal–diaphyseal ossi-
ica ion (An hony 1988).
Roos ees we e loca ed using s anda d adio eleme ic ech-
niques. We a ached LB-2 adio ansmi e s (0.46 g; Holohil Sys ems
L d., Woodlawn, On a io) o 11 ed ba s and 5 Seminole ba s. T ans-
mi e s we e a ached be ween he scapulae o he ba s wi h Skin
Bond® su gical adhesi e (P ize Hospi al P oduc s G oup, Inc.,
La go, Flo ida). The u was no clipped p io o a achmen o he
ansmi e s. T ansmi e load was less han 5% o he animal’s mass
(Ald idge and B igham 1988). Radio- agged ba s we e loca ed du ing
he day using Ad anced Teleme y Sys em R2000 ecei e s (Isand i,
Minneso a) and h ee-elemen Yagi an ennas. When oos ees could
no be eliably iden i ied using adio eleme y, oos loca ions we e
e i ied using binocula s du ing he day o by wa ching he oos o
eme ging ba s a dusk. We conduc ed exi coun s on 24 oos ees o
ed ba s and 16 oos ees o Seminole ba s. Fo oos ees on which
exi coun s we e conduc ed, 23 (96%) and 14 (87%) used by ed ba s
and Seminole ba s, espec i ely, we e co ec ly iden i ied using adio-
eleme y. In he h ee cases whe e he oos ee was inco ec ly
iden i ied using adio eleme y, he ac ual oos ee was wi hin 2 m.
In all h ee cases, he oos ee iden i ied using adio eleme y di -
e ed by less han 3 m in heigh and 15 cm in diame e om he ac ual
oos ee, and was o he same species as he ac ual oos ee. All ba s
we e acked as long as he ansmi e emained ope a ional and a -
ached o he ba . A e age ansmi e li e was 14 days. T ansmi e s
emained a ached o he ba s o an a e age o 8 days.
A e a oos ee was loca ed i was ma ked wi h lagging. Be-
ween 2 and 14 days a e he oos ing si e was loca ed, quan i a i e
measu emen s we e aken on he oos ee and he ege a ion su -
ounding i . A 0.04-ha sampling plo ( adius 11.35 m) was es ablished
a ound he oos ee. The diame e a b eas heigh (DBH), heigh ,
and species o all o e s o y ees (DBH >9 cm) wi hin he plo we e
measu ed. The species and s em numbe o all ees in he unde s o y
we e eco ded. Measu emen s aken on he oos ee included DBH,
o al heigh , and heigh o he base o he li e c own. All heigh s we e
measu ed using a clinome e . Pe cen canopy closu e was measu ed
o each plo using a sphe ical densiome e (Fo es Densiome e s,
A ling on, Vi ginia). We measu ed he densi y o he canopy 2 m
om he base o he oos ee on he side o he ee on which he ba
was oos ing. Measu emen s we e aken in he ou ca dinal di ec ions
and he mean densi y was eco ded. The aspec o he oos and he
heigh o he ba in he ee we e also eco ded. We measu ed aspec
using a compass and co ec ed o ue no h. Shannon’s di e si y
index and Pielou’s measu e we e calcula ed o bo h he o e s o y and
unde s o y ege a ional communi ies su ounding he oos s (Pielou
1966). Measu es o basal a ea we e used in calcula ing all o e s o y
indices. Unde s o y calcula ions a e based on he numbe o s ems pe
0.04-ha sampling plo .
The heigh s and DBHs o oos ees used by ed and Seminole
ba s we e compa ed wi h he a e age heigh and diame e o all o e -
s o y ees wi hin he sampling plo s. The a e age heigh and DBH o
all o e s o y ees su ounding he oos ee wi hin each 0.04-ha
sample plo we e calcula ed. The a e age heigh and DBH o all ees
su ounding he oos ees o ed ba s we e compa ed wi h he a e -
age heigh and DBH o all ees used as day oos s by ed ba s. The
a e age heigh and diame e we e also calcula ed o all ees su -
ounding he oos ees o Seminole ba s and he same compa isons
we e made. The use o each ee species as a day oos was compa ed
wi h he abundance o each species wi hin he sample plo s.
We de e mined he uni e sal ans e se me ca o (UTM) coo di-
na es o all oos ees using a T imble Pa h inde global posi ioning
sys em (GPS). Final coo dina es we e ob ained using di e en ial co -
ec ion. We calcula ed he oos a ea o i e adul ed ba s and ou
adul Seminole ba s using he minimal con ex polygon me hod and
he p og am calhome (Kie e al. 1996). The oos ing a ea calcula ed
o each indi idual was he a ea o a minimal con ex polygon ha
included all oos ees used by he indi idual while i was being
acked. Roos ing a eas we e compa ed be ween adul ed ba s and
Seminole ba s using a wo-sample es (SAS Ins i u e Inc. 1990).
We es ed o di e ences in all oos cha ac e is ics be ween oos s
loca ed on Sapelo Island and a he Sa annah Ri e Si e using a wo-
sample es . No signi ican di e ences we e de ec ed, he e o e da a
collec ed a bo h si es we e pooled o all analyses.
We compa ed 16 cha ac e is ics o day- oos si es be ween ed
ba s and Seminole ba s by means o a wo-sample es . The aspec s
o oos s selec ed by ed and Seminole ba s we e compa ed wi h
hose expec ed on he assump ion o a andom dis ibu ion, using a
3M.A. Menzel, D.M. K ishon, T.C. Ca e , and J. Lae m. No es
on ee oos cha ac e is ics o he no he n yellow ba (Lasiu us
in e medius), he Seminole ba (L. seminolus), he e ening ba
(Nyc iceius hume alis), and he eas e n pipis elle (Pipis ellus
sub la us).
Menzel e al. 631
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likelihood- a io es (Sokal and Rohl 1969). To examine wi hin-s and
oos selec ion, we compa ed he heigh s and DBHs o oos ees used
by ed and Seminole ba s wi h he a e age heigh and DBH o su -
ounding ees using a wo-sample es . The numbe o imes a ee
o a pa icula species was used as a oos was compa ed wi h he
abundance o ha species wi hin he sampling plo s, using a χ2 es .
Resul s
We acked 11 ed ba s and 5 Seminole ba s o hei day oos s.
The 64 oos ees o ed ba s we e used on 74 nigh s, he 34
oos ees o Seminole ba s on 57 nigh s. A e age oos -si e
ideli y o ed and Seminole ba s was 1.2 and 1.7 nigh s pe
oos ee, espec i ely.
Radio ansmi e s we e placed only on Seminole ba s cap u ed
a he sou h end o Sapelo Island. Al hough o e hal (54%)
o he s udy a ea on Sapelo Island consis ed o ha dwood-
domina ed communi ies, 25 oos s o Seminole ba s (88%)
we e loca ed in communi ies domina ed by pines. While he
Sa annah Ri e Si e s udy a ea con ained p ima ily pine-
domina ed communi ies (74%), 55 oos ees o ed ba s
(86%) we e loca ed in ha dwood communi ies. A he Sa an-
nah Ri e Si e, all nine (100%) o he oos s o Seminole ba s
we e loca ed in pine-domina ed communi ies. Ten o he 16
oos -si e cha ac e is ics examined di e ed signi ican ly be-
ween ed and Seminole ba s (Table 1). The 0.04-ha sampling
plo s su ounding he oos s o ed ba s con ained mo e o e -
s o y ees (mean ±SE =23.90 ±0.89) han he plo s su ound-
ing oos s o Seminole ba s (17.94 ±1.57). The o e s o y
su ounding he oos s o Seminole ba s (21.1 ±0.7 m) was
signi ican ly highe han ha su ounding he oos s o ed ba s
(17.3 ±0.5 m). Roos - ee diame e s also di e ed be ween he
wo species. The a e age DBH o ed ba oos ees (38.0 ±
2.2 cm) was signi ican ly smalle han ha o Seminole ba
oos ees (43.6 ±1.7 cm). Bo h he o e s o y and he unde -
s o y su ounding he oos s o ed ba s showed g ea e ich-
ness and di e si y han hose su ounding he oos s o Semi-
nole ba s (Table 1). The pe cen ages o coni e s in he o e -
s o y also di e ed signi ican ly be ween he oos s o ed and
Seminole ba s. On a e age, 20% o o e s o y ees su ound-
ing he oos s o ed ba s we e coni e s. In con as , he o e -
s o y su ounding he oos s o Seminole ba s consis ed, on
a e age, o 83% coni e s. Pe cen canopy closu e su ounding
he oos s o ed ba s (92.3 ±0.8%) was signi ican ly highe
han a ound he oos s o Seminole ba s (68.5 ±3.7%).
Red ba s we e ound oos ing in 18 species o ees (Table 2).
Swee gums (Liquidambe s y aci lua), black gums (Nyssa syl-
a ica), whi e oaks (Que cus alba), lau el-lea ed oaks
(Q. lau i olia), and wa e -oaks (Q. nig a) we e commonly
used. The species composi ion o oos ees di e ed signi i-
can ly om he composi ion expec ed in ha dwood habi a s,
assuming andom selec ion o ee species wi hin he oos ing
a ea (G=29.25, P=0.03). Red maples (Ace ub um) and
loblolly pines commonly occu ed wi hin he oos plo s; how-
e e , each species was used only once as a day oos . Whi e
oaks and wa e -oaks occu ed in equen ly wi hin he oos
plo , bu bo h species we e commonly used as day oos s
(Table 2). Bo h DBHs and heigh s o oos ees used by ed
ba s we e signi ican ly g ea e han he a e age DBH and
heigh o su ounding ees (Table 3). The oos aspec s se-
lec ed did no di e signi ican ly om a andom dis ibu ion.
Seminole ba s we e ound oos ing in i e species o ees
(Table 2). Loblolly pines, slash pines, and longlea pines we e
commonly used. The species composi ion o oos ees did no
Roos -si e cha ac e is ic L. bo ealis L. seminolus P
No. o ees in unde s o y 42.0±3.92 45.6±9.03 0.7161
No. o ees in o e s o y 23.9±0.89 17.9±1.57 0.0006
Heigh o o e s o y (m) 17.3±0.49 21.1±0.73 0.0001
Basal a ea o o e s o y
(m2/ha) 35.3±2.17 35.2±2.36 0.9767
Richness o unde s o y 6.9±0.33 3.7±0.39 0.0001
Di e si y o unde s o y 1.4±0.06 0.8±0.09 0.0001
E enness o unde s o y 0.7±0.02 0.6±0.05 0.0760
Richness o o e s o y 6.5±0.29 3.1±0.32 0.0001
Di e si y o o e s o y 1.3±0.04 0.5±0.90 0.0001
E enness o o e s o y 0.7±0.02 0.4±0.06 0.0001
Diame e o oos ee (cm) 38.0±2.24 43.6±1.69 0.0510
Heigh o oos ee (m) 25.0±0.99 27.8±1.03 0.0762
Roos heigh (m) 15.3±1.87 16.3±1.03 0.6432
Pe cen canopy closu e 92.3±0.76 68.5±3.68 0.0001
Pe cen age o snags in
o e s o y 2.6±0.01 2.0±0.01 0.6097
Pe cen age o coni e s in
o e s o y 19.7±0.03 82.9±0.04 0.0001
No e: Values a e gi en as he mean ±SE.
Table 1. Compa ison o 16 oos -si e cha ac e is ics be ween
Lasiu us bo ealis and L. seminolus.
L. bo ealis L. seminolus
T ee species Roos ing Random Roos ing Random
Ace leucode me 1.6 0.1 0 0
A. ub um 1.6 4.7 0 0
Ca ya glab a 1.6 0.9 0 0
Liquidambe
s y aci lua 24.6 25.2 0 0
Li iodend on
ulipi e a 4.9 6.0 0 0
Nyssa aqua ica 3.4 5.6 0 0
N. syl a ica 8.3 7.0 0 0
Pla anus occiden alis 1.6 0.4 0 0
Que cus alba 9.8 4.8 0 0
Q. du andii 3.4 0.2 0 0
Q. alca a 1.6 0.6 0 0
Q. michauxii 1.6 1.4 0 0
Q. lau i olia 8.3 9.6 0 0
Q. nig a 21.3 15.1 0 0
Q. s ella a 1.6 1.1 0 0
Q. i giniana 0 0 5.6 3.0
Ulmus ame icana 1.6 2.3 0 0
Pinus palus is 1.6 0.8 13.9 10.7
P. aeda 1.6 14.2 30.5 25.4
P. ellio ii 0 0 47.2 55.6
P. se o ina 0 0 2.8 5.3
Table 2. P opo ion o each species o ee used o oos ing
compa ed wi h he abundance o ha species wi hin he 0.04-ha
sampling plo su ounding he oos s o Lasiu us bo ealis and
L. seminolus.
Can. J. Zool. Vol. 76, 1998
632
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di e signi ican ly om he expec ed composi ion wi hin
pine-domina ed habi a s. Like ed ba s, Seminole ba s oos ed
in ees o g ea e heigh s and DBHs han he a e age alues
o ees wi hin he oos plo (Table 3). Roos ing si es we e
selec ed andomly wi h espec o aspec .
Roos ing a eas di e ed signi ican ly be ween ed and Semi-
nole ba s. Bo h species mo ed o new oos ees o en. Red
ba s and Seminole ba s spen an a e age o 1.2 and 1.7 days
on one oos ee, espec i ely. While indi iduals o bo h spe-
cies o en mo ed o new oos ees, adul ed ba s selec ed
oos s wi hin a signi ican ly la ge a ea (2.6 ±0.6 ha) han
adul Seminole ba s (0.2 ±0.1 ha; P=0.01).
Discussion
The di e ences be ween he oos ing si es o ed and Seminole
ba s we e ela ed di ec ly o he di e en oos ing habi a s used
by he wo species. The oos s o ed ba s we e loca ed bo h in
pine – mixed ha dwood communi ies domina ed by swee -
gums, wa e -oaks, and loblolly pines and in bo omland ha d-
wood swamps domina ed by co on-gum (Nyssa aqua ica) and
bald cyp ess (Taxodium dis ichum). The oos s o Seminole
ba s we e always loca ed in o es communi ies domina ed by
pines. Bo h on Sapleo Island and a he Sa annah Ri e Si e,
pine-domina ed communi ies a e in ensi ely managed o he
p oduc ion o o es p oduc s. S ands o pines we e ypically
composed o a single ee species and all indi idual ees we e
o app oxima ely he same age and size. The unde s o y com-
muni ies su ounding he oos s o Seminole ba s we e man-
aged wi h he use o p esc ibed i es, esul ing in a open
unde s o y o sh ub communi ies domina ed by a ew i e-
adap ed species. The unde s o y communi ies su ounding he
oos s o Seminole ba s we e less di e se han hose su ound-
ing he oos s o ed ba s. The pine – mixed ha dwood commu-
ni ies and bo omland ha dwood swamps whe e ed ba s
oos ed we e managed much less in ensi ely. These s ands
we e no hinned. Because ed ba s oos ed in less in ensi ely
managed s ands, hei oos s we e su ounded by mo e o e -
s o y ees han hose o Seminole ba s. Since he e we e mo e
ees in he o e s o y, canopy densi y su ounding he oos s o
ed ba s was g ea e han ha su ounding he oos s o Semi-
nole ba s. Because o he une en-aged o e s o y in he less
in ensi ely managed s ands, a e age o e s o y heigh s we e
lowe and a e age oos - ee diame e s smalle o ed ba s
han o Seminole ba s.
Excep o hei heigh in he ee, he oos s o ed ba s we e
simila o hose desc ibed in he li e a u e. We ound ed ba s
oos ing a an a e age heigh o 15.3 m, highe han hose
ound by Mum o d (1973; 2.6 m), Koon z and Da is (1991;
1.5 m), and McClu e (1942; 1.6–12.5 m). P e ious in es iga-
ions in o he oos ing habi s o ed ba s in ol ed isually lo-
ca ing he oos s. Ba s oos ing lowe in he ee may be loca ed
mo e easily by isual sea ches han hose oos ing high in he
canopy (Mum o d 1973). Because we loca ed ou oos s by
means o adio eleme y, ou es ima es o oos ing heigh s a e
p obably mo e ep esen a i e han p e ious es ima es. Like
McClu e (1942) and Cons an ine (1966), we ound ed ba s
oos ing in many species o ees. Al hough McClu e (1942)
compa ed he p opo ions o oos s loca ed in di e en species
o ees wi h he ela i e abundance o each ee species, he
did no de e mine i any ee species we e used mo e o en han
expec ed. We ound ha ed ba s p e e ed whi e oaks and
wa e -oaks and did no oos in ed maples o loblolly pines as
o en as expec ed.
The oos s o Seminole ba s ha we ound we e unlike hose
p e iously desc ibed. Cons an ine (1958) ound Seminole ba s
oos ing a an a e age heigh o 2.5 m, bu we ound hem
oos ing a an a e age heigh o 16.3 m. Roos s o Seminole
ba s we e commonly loca ed in Spanish moss (Ba bou and
Da is 1969; Cons an ine 1958; Ha pe 1927; Jennings 1958;
Wilkins 1987). Al hough Seminole ba s o age in pine-domi-
na ed communi ies (Ha pe 1927; I ey 1959; Jennings 1958;
Moo e 1949; Zinn 1977), we could ind no epo s o Seminole
ba s oos ing in pines. The Seminole ba s we acked almos
always oos ed in pines. Cons an ine (1958) sea ched o
oos s o Seminole ba s in Spanish moss om Feb ua y
h ough June. He loca ed ba s in e e y mon h excep May and
June. Jennings (1958) collec ed oos ing Seminole ba s om
Spanish moss h oughou he win e , bu ailed o collec any
du ing July, Augus , o Sep embe . Jennings examined Span-
ish moss o ba s h oughou he yea , bu ew li e s o Semi-
nole ba s we e ound. Thus, du ing he pe iod o pa u i ion
and lac a ion, Seminole ba s may commonly oos in pines.
Many species o ba s use di e en ypes o oos du ing he
yea in esponse o di e ing physiological demands
(e.g., hibe na ion, p egnancy, lac a ion; Ba bou and Da is
1969). Thus, he di e ences be ween he oos s we loca ed and
hose p e iously desc ibed likely ela e o he ime o he yea .
Cons an ine (1958, 1966) sugges ed ha ed ba s and Semi-
nole ba s selec oos s on he sou hwes e n and sou he n side
o oos ees, espec i ely. We ound ha he oos s o bo h
species we e selec ed andomly wi h espec o aspec .
Compa isons be ween he size o ees used as oos s by ed
ba s o Seminole ba s and he a e age size o su ounding ees
in he o e s o y a e lacking. Howe e , Vonho and Ba clay
(1996) made gene al compa isons be ween ees used as oos s
by ou species o ba s in B i ish Columbia and ees selec ed
andomly om he o e s o y. The ees used as oos s we e
signi ican ly alle han andomly selec ed ees. Roos ees
selec ed by ou species o ba s in Tasmania had la ge diame-
e s han andomly selec ed ees (Taylo and Sa a 1988). We
ound ha oos ees used by ed and Seminole ba s we e
L. bo ealis L. seminolus
Roos Random PRoos Random P
DBH (cm) 37.75±2.22 22.53±0.70 0.0001 42.08±1.84 30.18±1.27 0.0001
Heigh (m) 24.85±1.00 16.96±0.48 0.0001 26.51±1.07 20.72±0.72 0.0001
No e: Values a e gi en as he mean ±SE.
Table 3. Compa ison o he diame e s and heigh s o ees used as oos s by Lasiu us bo ealis and L. seminolus
wi h he a e age diame e and heigh o all o e s o y ees wi hin he 0.04-ha plo s su ounding he oos s.
Menzel e al. 633
© 1998 NRC Canada
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signi ican ly alle and had la ge diame e s han su ounding
ees. Ou esul s suppo he indings o Vonho and Ba clay
(1996) and Taylo and Sa a (1988).
Adul ed ba s possibly selec oos s om la ge a eas han
adul Seminole ba s because o he ela i e a ailabili y o he
habi a ypes in which hey p e e ed o oos . On bo h o ou
s udy si es, pine – mixed ha dwood communi ies we e mo e
abundan han s ands composed only o pine. The oos s o
Seminole ba s we e ound in islands o pine monocul u e su -
ounded by pine – mixed oak communi ies. Seminole ba s had
o ei he es ic shi s in oos loca ion o o he ees wi h he
island o make majo shi s o di e en islands. Because he
ype o oos ing habi a p e e ed by ed ba s was ai ly ubiq-
ui ous, hey we e ee o selec oos s om la ge a eas han
Seminole ba s.
These da a on he oos ing habi s o Seminole ba s aise new
ques ions conce ning he e ec s o summe imbe -ha es ing
ope a ions in he sou heas e n Uni ed S a es on he ep oduc-
i e success o his species. Fu u e s udies in es iga ing he
esponse o Seminole ba s o imbe ha es ing in upland pine
s ands a e needed.
Acknowledgmen s
We hank E.B. Higgins, S.B. Cas lebe y, N. an de Maa h,
N. Hicks, M. Wi che , R. Hendi, C. Moo eman, and T.S.
McCay o ield assis ance, M. Da alos o assis ance wi h
GPS equipmen and A c In o, and he Sa annah Ri e Ecology
Labo a o y o p o essional suppo and he use o eleme y
equipmen . We also hank R.J. Wa en, J. Kilgo, and W.M.
Fo d o edi o ial ad ice. This p ojec was unded by he
Uni ed S a es Fo es Se ice.
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