P oceedings o he Indiana Academy o Science J27
(1992) Volume 101 p. 127-134
SEASONAL ACTIVITY OF BATS
AT COPPERHEAD CAVE
John O. Whi ake , J . and Leslie J. Rissle
Depa men o Li e Sciences
Indiana S a e Uni e si y
Te e Hau e, Indiana 47809
Abs ac : Coppe head Ca e is an abandoned mine in Ve million Coun y, Indiana, se ing as bo h
ahibe naculum and aswa ming a ea o no he n ba s (Myo is sep en ionalis), li le b own ba s
(M. luci ugus), and eas e n pipis elles {Pipis ellus sub la us). Adouble- ame ha p ap was used
o documen ba ac i i y a he mou h o he mine on aweekly basis om he all o 1989 o he
sp ing o 1991. Mos o he apped ba s we e banded and eleased. Li le b own ba s o bo h
sexes eme ged g adually om hibe na ion be ween 14 Ma ch and 19 May 1990 and be ween 15
Ma ch and 1May 1991. In con as , no he n ba s eme ged in much la ge numbe s be ween 9
Ma ch and 27 Ap il 1990 and be ween 21 Feb ua y and 22 Ap il 1991. The mine se ed as a
swa ming a ea om he end o July o he end o Oc obe o bo h species o Myo is and h ough
he i s week o Oc obe o he pipis elles. The pipis elles exhibi ed almos no win e ac i i y
( ligh ). Howe e , he li le b own ba s and he no he n ba s we e ac i e a he mine en ance
h oughou he win e . Tu no e in he ba popula ions o his mine is qui e high, as indica ed by
he low numbe o banded ba s eco e ed: 336 o 1733 (19.4%) indi iduals o Myo is luci ugus;
174 o 2090 (8.3%) indi iduals o M. sep en ionalis; and 100 o 546 (18.3%) indi iduals o
Pipis ellus sub la us.
INTRODUCTION
Coppe head Ca e is an abandoned clay mine in ablu 1km wes o he Wabash
Ri e and 1.5 km wes o Mon ezuma in Ve million Coun y, Indiana. The mine is abou
120 km no hwes o he edge o he unglacia ed ca e coun y o sou he n Indiana, and i
ep esen s an a i icial ca e in an a ea whe e no na u al ca es exis . We we e in o med
ha he mine se ed as ahibe naculum o ba s, and ini ial moni o ing indica ed ha he
mine was used in o he seasons as well.
Regula moni o ing o he mine en ance was ini ia ed o de e mine:
1. The species, numbe , and sex o he ba s using he mine;
2. The seasonal use o he mine;
3. The uses o which he mine was pu ;
4. The da es o en ance in o and exi om hibe na ion;
5. The size o he ba popula ion; and
6. Whe he o no as able ba popula ion used he mine o ex ended pe iods
o ime, o i signi ican popula ion u no e occu ed.
MATERIALS AND METHODS
Adouble ame ha p ap was used o sample ba s lying a he en ance o he mine.
Moni o ing s a ed in Sep embe 1989 and was o be e mina ed o he season on 30 Oc-
obe , because we assumed ha ba ac i i y would essen ially cease by ha ime. How-
e e , o e i y ha li le o no win e ac i i y occu ed, he ap was placed a he mine
128 ZooLOGY:Whi ake and Rissie Vol. 101 (1992)
Table 1.Summa y o in o ma ion on he ba s aken in a ap a he en ance o Coppe head
Ca e.
Da e M. 'uci ugus M. sep en ionalis Pipis ellus
male emale male emale male emale
1989
Sep 19 110 14 41 27 43
22 100 10 310 6
26 62 19 710 34
30 67 710 18 25
Oc 340 11 117
10 24 11 16 21 23 15
13 21 564 38 32
17 31 571 1
20 92421
24 72 41 12 1
28 8 3 28 82
30 7415 6
1990
Jan 16 9312
23 3412
27 2 2 2
Feb 311
8736511
17 1
23
Ma 31521
93112 41
14 3 3 61 6
17 315 1
24 121 31
31 570 44
Ap 432 32
71437 62 1
11 4510 10
17 37919
27 77912 8
May 6213
94124
12 2211
19 244
26 13 3
Vol. 101(1992) Indiana Academy o Sen2NCE 129
Da e M. ihici iigus Msep en ionalis Pipis i ellus
male emale male emale male emale
June 23 3
15 278 21
23 175 1
29 2 35 2
30 357 1
July 1232 1
2321 2
36119 3
4320 10
5717 6
6671
10 117
17 219
25 815 8 1
31 8331 11 5 2
Aug 79731
15 22 23 66 56 63
20 42 37 35 52 22
25 47 46 82 89 33 15
Sep 244 25 48 55 45
947 12 12 34 84
13 40 918 17 26
21 53 17 15 28 1
26 36 933 49 49
Oc 512 625 37 13
12 10 112 81
19 11 110 1
29 2113 6
No 84I21
15 1242
20 1348
30 142
Dec 8112
16 123
21 114
1991
Jan 2 112
10 332
18 2
25 21
130 ZooLOGY:Whi ake and Rissle Vol. 101 (1992)
Da e M.iluci ugus M. sep en ionalis Pipis ellus
male emale male emale male emale
Feb 2312
81
17 312
21 1310
28 34713
Ma 3165 149 7684 86
7112 2
15 410 713
22 212 5
29 15 5632
Ap 51130 10
22 4411 811
May 12288
9145
en ance on 21 No embe 1989, le in place, and checked daily h ough ea ly Decembe .
A e ea ly Decembe , he ap was checked spo adically. Th ough 9Janua y 1990, a o al
o 63 ba s was aken in his un ended ap: 22 in No embe , 22 in Decembe , and 19 in
Janua y. Included we e 32 li le b own ba s, 29 no he n ba s, and 2pipis elles. Much
mo e win e ac i i y occu ed han had been an icipa ed. The e o e, s a ing on 16 Janua y
1990, he ap was ended app oxima ely one nigh pe week un il 9May 1991 (Table 1).
In addi ion, an assessmen o he ba s hibe na ing in he mine was made on 23 Feb ua y
1988, and simila assessmen s we e made on 14 Janua y 1989, 27/29 Decembe 1989, and
3Ma ch 1991 (Table 2). Nea ly all he ba s aken in he ap and nea ly all o hose ound
in hibe na ion du ing he la e wo assessmen s we e banded (plas ic numbe ed bands om
Na ional Band and Tag Co, Newpo , Ken ucky). The bands emained on he ba s qui e
well, bu he numbe s o en become di icul o ead wi hin ayea o wo. Fou housand,
h ee hund ed six y-nine ba s we e banded, and 610 we e ecap u ed a Coppe head Ca e.
RESULTS
Ba s use his mine yea ound (Table 1), al hough he amoun o use a ies season-
ally. Dec eased ac i i y was encoun e ed a he mine en ance om No embe h ough
Feb ua y and om ea ly in May h ough he i s week in June. The majo i y o ba s en-
coun e ed belonged o h ee species: he li le b own ba , Myo is luci ugus, he no he n
ba , Myo is sep en ionalis, and he pipis elle, Pipis ellus sub la us. All h ee species
swa m he e in he all, p esumably o ma ing pu poses, and all h ee also hibe na e in he
mine. A ew ba s o wo o he species we e also aken: se en Indiana ba s {Myo is sodalis)
and i e big b own ba s (Ep esicus uscus). These ba s will no be discussed u he .
O pa icula in e es was he ac ha no he n and li le b own ba s we e ac i e all
win e a his mine. Inc eased ac i i y occu ed on wa me nigh s (Whi ake and Rissle ,
1992). The e was li le win e ac i i y by pipis elles. The ba s en e ing and lea ing he
Vol. 101 (1992) Indiana Academy o Science 131
Table 2. Ba s in hibe na ion in Coppe head Ca e.
Da e M. luci ugus M. sep en ionalis Pipis ellus
male emale male emale male emale
Feb. 23, 1988 111* 201*
Jan. 14, 1989 106* 113*
Dec. 27/29, 1989 85 48 47 45
Ma ch 3, 1991 165 149 7684 86
Ba s we e coun ed wi hou emo ing hem om he wall; hus, in o ma ion on sex is no
a ailable.
mine in midwin e do no eed e en on wa m nigh s, when insec s a e lying (Whi ake
and Rissle , 1992). Some o he win e ligh s may e lec mo emen s be ween hibe nacula.
Howe e , some o he lying may occu because he ba s a e simply lying ou side a he
han inside du ing no mal win e awakenings om hibe na ion (Cock um, 1956; Menake ,
1964; Mum o d, 1958). The o al numbe o ba s aken in 1990 and 1991 du ing he win-
e apping sessions included: 27 no he n ba s and 10 li le b own ba s in Decembe ; 15
no he n ba s and 20 li le b own ba s in Janua y; and 47 no he n ba s and 30 li le b own
ba s in Feb ua y. Only h ee pipis elles we e aken du ing he win e apping sessions.
Myo is luci ugus. Myo is luci ugus usually hibe na es in ca es, bu his mine p o-
ides aca e-like en i onmen in an a ea wi h no ca es. The mine se ed as ahibe nacu-
lum o he ollowing minimum numbe o li le b own ba s on he ollowing da es: 111on
23 Feb ua y 1988; 72 on 14 Janua y 1989; 133 on 27/29 Decembe 1989; and 314 on 3
Ma ch 1991.
No majo peak o sp ing eme gence ac i i y by li le b own ba s was obse ed in ei-
he 1990 o 1991. The ela i ely small numbe o ba s p esen p obably eme ged a he
g adually om Ma ch h ough May. These da a co ela e ai ly well wi h da a on he buildup
in ama e ni y colony o his species a B azil, Clay Coun y, Indiana. A B azil, eme -
gence s a ed g adually in la e Ma ch and was la gely comple e by la e Ap il (Ronald Da is,
unpublished da a).
Coppe head Ca e did no se e as amajo bachelo qua e s o his species in he
summe , as he e was ela i ely li le ac i i y du ing June and July 1990. Be ween wo o
eigh ba s, including only h ee emales, we e aken each nigh om 1June o 25 July. On
31 July, eigh males and h ee emales we e aken, ollowed by se en emales and nine
males on 7Augus 1990. Thus, he end o July and ea ly Augus appea o ma k he be-
ginning o swa ming o his species. These da es co ela e qui e well wi h adec ease in
he summe popula ion in he colony a B azil in Clay Coun y, whe e he numbe o ba s
p esen d opped o d as ically beginning in ea ly Augus . Swa ming ape ed o in Oc o-
be , wi h swa ming by emales dec easing soone han ha by males (10 Oc obe e sus
17 Oc obe in 1990; 5Oc obe e sus 19 Oc obe in 1991). Appa en ly, he ba s mo e
g adually o hei swa ming a eas and hen g adually in o hibe na ion. Some o he ba s
hibe na e a Coppe head Ca e, bu mos mo e elsewhe e, as indica ed by he low numbe
o banded ba s ound in hibe na ion. P io o 3Ma ch 1991, a o al o 1470 li le b own
ba s had been banded in o a he en ance o Coppe head Ca e, whe eas on 3Ma ch, only
72 banded li le b own ba s we e ound hibe na ing he e. This numbe ep esen s only
4.9% o he o al numbe o ba s banded. Se e al o he ba s banded a Coppe head Ca e
we e ound hibe na ing du ing he win e o 1990-91 abou 100 km o he sou hwes in
132 ZooLOGY:Whi ake and Rissle Vol. 101 (1992)
Table 3. The o al numbe o ba s banded a Coppe head Ca e.
Species Male Female Unknown
M. luci ugus 1234 491 8
M. sep en ionalis 1431 652 7
Pipis ellus 325 219 2
Ep esicus 41
M. sodalis 43
Ray's and Cly y Ca es in he Law ence Coun y a ea.
Myo is sep en ionalis. Coppe head Ca e appa en ly se es as ahibe naculum o
no he n ba s, al hough no many indi iduals we e ound in hibe na ion. Th ee lines o
e idence indica e ha he mine is ahibe naculum: he (1) numbe o no he n ba s ha
en e ed he ca e in all; he (2) numbe ha eme ged in win e and sp ing; and he (3) ba s
ound hibe na ing in iny c acks in win e . The nume ous c acks in his mine mus se e
as win e qua e s o hund eds o no he n ba s, bu inding hem was e y di icul in hese
c acks. On ela i ely wa m sp ing nigh s, when no he n ba s we e eme ging om he mine,
nume ous indi iduals we e ound exi ing om he c acks (e.g., wel e we e obse ed on
15 Ma ch 1991).
No he n ba s eme ge om hibe na ion ea ly in he season. La ge numbe s exi ed om
he mine on nigh s when he empe a u e app oached 10° C om he second week o Ma ch
h ough Ap il each yea (Table 1). The modal exodus ime o males was app oxima ely
one week ahead o ha o emales.
An a emp was made o es ima e he numbe o no he n ba s using Coppe head Ca e
in sp ing. Sampling imes a e aged 7.5 hou s o 0.625 o he da k pe iod a ha ime o
yea . To de e mine i banded ba s migh be een e ing he mine o i ba ac i i y migh
ape o a e ou no mal sampling pe iod ended, ba s we e apped h oughou he nigh
on wo sepa a e occasions. The esul s indica ed ha ba ac i i y con inued all nigh and
ha indi iduals seen ea lie did no gene ally e u n o he mine a e ou no mal sampling
pe iod ended. The eme gence o no he n ba s in 1990 s a ed abou 9Ma ch and las ed
h ough 17 Ap il, a o al o 40 nigh s. Samples we e collec ed on en o hese nigh s. The
a e age numbe o no he n ba s aken pe nigh was 22.7 (4.2 ba s/0.625 =22.7 ba s; SD
=15.9, SE =5.3). Based on his numbe , an es ima ed 908 no he n ba s exi ed he mine
in sp ing (22.7 ba s/nigh x40 nigh s =908 ba s). This igu e is a en a i e, minimal es i-
ma e o he hibe na ing popula ion.
Du ing May and in o ea ly June 1990, ew no he n ba s we e ac i e a Coppe head
Ca e. Six we e aken in May, and h ee we e aken on 2June. Howe e , om 15 June o
5July, la ge numbe s o males bu only wo emales we e aken (a o al o 354 ba s was
aken in nine sessions). Ou i s in e p e a ion was ha he mine was se ing as abach-
elo colony. This would ha e been ou inal in e p e a ion, i we had sampled only once.
Howe e , i he males we e qua e ing he e, we should ha e had many ecap u es. Only
10.7% o he cap u es we e o banded ba s (38 ecap u es ou o 354 o al cap u es). O
he ecap u ed ba s, only 2.3% (eigh ) we e o iginally banded du ing he i s sampling pe-
Vol. 101 (1992) Indiana Academy o Science 133
iod. The da a sugges ha hese no he n ba s we e summe mig an s, al hough we do no
know whe e hey would be going o why. The la ge numbe o no he n ba s ha we e
p esen in his h ee week pe iod p ecludes hei being simply summe wande e s. We a e
unawa e o o he da a which include ex ended obse a ions on concen a ions o males in
summe .
Fall swa ming began a abou he same ime as i did o he li le b own ba . How-
e e , he onse was mo e ab up . Thi y-one males and 11 emales we e cap u ed on 3
1
July, a e only i e had been cap u ed six days p e iously. Majo swa ming ac i i y las ed
h ough he i s week in Oc obe . A e ha , he numbe o no he n ba s aken pe sample
emained low h ough Decembe . I was no possible o ell when all swa ming s opped,
because no he n ba s we e aken h oughou he win e .
Pipis ellus sub la us. The numbe s o pipis elles coun ed while hibe na ing in he
mine we e: 201 on 23 Feb ua y 1988; 113 on 14 Janua y 1989; 99 on 27/29 Decembe
1989; and 176 on 3Ma ch 1991. Pipis elles appa en ly mo e abou e y li le in hei
hibe naculum du ing win e . The wa e d ople s ha accumula e on hem a ha ime in-
dica e long- e m hibe na ion in con as o he equen awakenings cha ac e is ic o Myo is.
Mos pipis elles had le hibe na ion by he end o Ap il in 1990: none we e aken on 17
Ap il; 20 we e aken on 27 Ap il; and only 3we e aken on 6May. Howe e , a ew
pipis elle emales we e s ill lea ing he mine h ough he i s wo weeks in May. In 1991,
hey le la e : 16 we e aken on 1May, and 9we e aken on 9May. This ag ees wi h ou
unpublished da a indica ing he o ma ion o pipis elle ma e ni y colonies beginning in la e
Ap il and ex ending h ough much o May.
Small numbe s o male pipis elles we e aken om 4 July h ough 31July. The mine
may ha e se ed as amig a o y s opo e o hese ba s. Al e na i ely, hese males may
ha e ep esen ed ea ly swa ming indi iduals, al hough emales we e no included in swa m-
ing un il he end o July. The la ges numbe o swa ming indi iduals was encoun e ed on
25 Augus (33 males and 15 emales). Swa ming ac i i y occu ed h oughou all o Au-
gus and Sep embe and in o he i s week o Oc obe . A ha ime, he ba s ei he mo ed
elsewhe e o en e ed he ca e o hibe na e. A e 12 Oc obe 1990, no pipis elles en e ed
he ap un il 29 Ma ch 1991. In 1989-90, howe e , i e pipis elles we e aken on 13 Oc-
obe , i e mo e we e aken la e in Oc obe , and 13 we e aken be ween No embe and
Ap il.
CONCLUSION
Coppe head Ca e se es as aswa ming a ea and hibe naculum o li le b own ba s,
no he n ba s, and pipis elles. In addi ion, he mine may se e as amidsumme s opo e
o mig a ing no he n ba s and pipis elles.
LITERATURE CITED
A e y, M.I. 1985. Win e ac i i y by pipis elle ba s. J. Anim. Ecol. 54: 721-738.
B ack, V. and J.W. Twen e. 1985. The du a ion o he pe iod o hibe na ion o h ee species o espe ilionid
ba s. I. Field s udies. Can. J. Zool. 63: 2952-2954.
Cock um, E.L. 1956. Homing, mo emen s, and longe i y o ba s. J. Mammal. 37: 48-57.
Menake , M. 1964. The equency o spon aneous a ousal om hibe na ion in ba s. Na u e 203: 540-541.
Mum o d, R.E. 1958. Popula ion u no e in win e ing ba s in Indiana. J. Mammal. 39: 253-261.
Whi ake , J.O., J . and L.J. Rissle . 1992. Win e ac i i y o ba s a amine en ance in Ve million Coun y.
Indiana. Ame . Midi. Na u . 127: 52-59.
