ROOSTING?BEHAVIOR?AND?HABITAT?DYNAMICS?OFMALE?INDIANA?BATS?(MYOTIS?SODALIS)?FOLLOWING?A?LARGE?SCALE?NATURAL?DISTURBANCE? Except?where?reference?is?made?to?the?work?of?others,?the?work?described?in?this?disertation?is?my?own?or?was?done?in?colaborationwith?my?advisory?commitee.?This?disertationdoes?not?include?proprietary?or?clasified?information.? __________________________________________?Paul?Roger?Moosman,?Jr.? Certificate?of?Approval:? _______________________?_______________________?Robert?S.?Lishak? Troy?L.?Best,?Chair?Associate?Profesor? Profesor?Biological?Sciences? Biological?Sciences? _______________________?_______________________?Gary?R.?Hepp? Stephen?L.?McFarland?Profesor? Acting?Dean?Forestry?and?Wildlife?Sciences? Graduate?School ROOSTING?BEHAVIOR?AND?HABITAT?DYNAMICS?OFMALE?INDIANA?BATS?(MYOTIS?SODALIS)?FOLLOWING?A?LARGE?SCALE?NATURAL?DISTURBANCE? Paul?R.?Moosman,?Jr.? A?Disertation?Submited?to? the?Graduate?Faculty?of?Auburn?University? in?Partial?Fulfillment?of?the?Requirements?for?the? Degree?of?Doctor?of?Philosophy? Auburn,?Alabama?August?8,?2005 iii? VITA?Paul?Roger?Moosman,?Jr.?was?born?to?Paul?R.?Moosman,?Sr.?and?Jacqueline?T.? Moosman?in?London?Dery,?New?Hampshire,?22?August?1976.?While?growing?up?in?New?Hampshire,?he?developed?an?early?love?of?nature??spending?many?hours?catching? crayfishes,?frogs,?and?snakes?at?the?local?swamp,?or?fishing?with?his?father?and?uncles.?It?was?here?also,that?the?effects?of?urban?sprawl?and?development?left?an?early?impression? on?Paul.?At?age?14,the?Moosman?family?moved?to?the?smal?town?of?Brookneal,?Virginia,?where?there?was?litle?to?do?except?fish,?hunt,?or?spend?the?day?scouting?for? Native?American?artifacts.?After?finishing?high?schol?at?William?Campbell,?in?Naruna,?Paul?attended?the?Virginia?Military?Institute(VMI)thanks?to?anacademic?scholarship.? Life?at?VMI?was?a?struggle?to?balance?academics,?duties?to?the?rifle?team,?and?the?perpetual?demands?of?a?military?system.?However,?Paul?survived,?and?in?1998?left?VMI? with?many?close?friends?and?a?Bachelor?of?Science?degree?in?biology.?Imediately?after?graduating?from?VMI,?Paul?began?working?towardhis?M.S.?degree?in?biology,?at?Eastern? Kentucky?University(EKU),?in?Richmond.?While?at?EKU?he?conducted?field?research?on?bats?and?received?his?first?formal?training?in?subjects?that?he?loved,?such?as?aquatic,? marine,?and?terrestrial?ecology.?Paul?graduated?from?EKU?in?2001?and?went?on?to?complete?his?Ph.D.?disertation?at?Auburn?University,?under?the?direction?of?TroyL.?Best. iv? DISSERTATION?ABSTRACT? ROOSTING?BEHAVIOR?AND?HABITAT?DYNAMICS?OFMALE?INDIANA?BATS?(MYOTIS?SODALIS)?FOLLOWING?A?LARGE?SCALE?NATURAL?DISTURBANCE? Paul?Roger?Moosman,?Jr.?Doctor?of?Philosophy,?May?13,?2005?(M.S.,?Eastern?Kentucky?University,?2001)? (B.S.,?Virginia?Military?Institute,?1998)?95?Typed?Pages? Directed?by?Troy?L.?Best?Indiana?bats?(Myotis?sodalis)?have?been?the?subject?of?numerous?ecological? studies,?partly?because?they?are?endangered.?A?majority?of?research?has?focused?on?maternity?colonies?and?relatively?les?is?known?about?males.?Previous?research?identified? habitat?and?described?fidelity?of?a?population?of?male?Indiana?bats?at?South?Goldson?Cave,?Pulaski?Co.,?Kentucky.?Mortality?of?pine?trees?(Pinus)?folowing?a?large? infestation?of?southern?pine?beetles?(Dendroctonus?frontalis)?at?the?site?provided?an?opportunity?to?gain?insight?into?ecological?requirements?of?male?Indiana?bats,?and? determine?if?availability?of?dead?trees?influences?fidelity.?This?disertation1)?provides?a?sumary?of?the?conservation?and?life?history?of?Indiana?bats,?2)identifies?microhabitat? characteristics?ofmale?Indiana?bats?using?natural?roosts,?and?northern?long?eared?bats?(Myotis?septentrionalis)?using?bat?boxes,?3)?identifies?macrohabitat?variables?that v? influenced?selection?by?Indiana?bats,?4)determines?if?fidelity?changed?folowing?the?infestation?ofsouthernpine?beetles,?and?5)?estimates?how?long?roosts?may?remain?suitable? for?use?by?bats.?Most?roosts?were?under?exfoliating?bark?ofdead?pine?trees,?ca.?9?m?from?the?ground,?with?sunny?or?partially?shaded?conditions.?Bat?boxes?were?used?infrequently? by?northern?long?eared?bats?(7?of?46?boxes),?primarily?during?summer.?Inadequate?temperature?regimes?may?have?limited?use?of?bat?boxes?to?summer.?Habitats?selected?by? Indiana?bats?occurred?in?stands?1)?dominated?by?pines?with?few??other??trees?present,?and?2)?with?greater?mean?diameters?at?breast?height,?basal?area,?abundant?dead?hardwoods,? and?less?abundant?living?hardwoods.?Use?ofcore?areas?over?time?was?influenced?by?local?availability?of?dead?trees.?Bats?used?pine?trees?as?they?became?available?folowing?the? infestation?of?southern?pine?beetles.?Fidelity?increased?slightly?during?2001?2003,?but?was?statisticallysimilar?to?data?reported?priorto?the?infestation.?Roosts?decayed?over? time??most?pines?were?predicted?to?become?unsuitable?2?7?years?from?the?date?they?were?used?by?bats,?whereas?hardwoods?were?predicted?to?remain?suitable?indefinitely.?Used? habitats?likely?provided?warmtemperatures?that?aided?in?thermoregulation.?Abundant?dead?hardwood?trees?in?habitats?that?were?used?by?Indiana?bats?suggeststhat?1)?bats?use? areas?with?many?potential?alternate?roosts,?and?2)?hardwoods?may?have?ben?used?frequently?before?natural?disturbances?killed?large?numbers?of?pines.?Although?male? Indiana?bats?used?ephemeral?rosts?created?by?natural?disturbance,?bats?showed?fidelity?to?core?areas?as?wel?as?individual?trees.?Importance?of?natural?disturbances?to?bats?may? depend?on?time?intervals?between?disturbances. vi? ACKNOWLEDGMENTS?This?project?would?have?been?extremely?difficult?if?not?for?the?support?ofmany? people.?My?wife,?Deanne?Moosman,?deserves?more?credit?than?I?wil?ever?be?able?to?show.?More?than?just?helping?me?track?bats?through?rain,?or?cary?bat?boxes?and?concrete? through?a?sea?of?thorns?and?fallen?logs,?she?truly?was?my?light?at?the?end?of?this?tunnel.?I?also?thank?Joe?Metzmeier?for?his?hospitality??all?the?hours?spent?sitting?in?front?of?those? fires?were?wel?spent.?Additionally,?I?am?grateful?for?the?help?and?friendship?of?Mark?Gumbert?and?Brainard?Palmer?Ball,?Jr.?who?donated?sweat?and?blod?on?many?occasions? to?help?capture?and?track?bats.?Also,?Ithank?Joy?O?Keefe?for?her?companionship?in?the?field?and?advice?with?Geographic?Information?Systems.? Numerous?others?deserve?mention?for?help?they?provided?in?the?form?of?assistance?in?the?field,?logistical?support,?or?discussion?of?ideas,?including?John?MacGregor,?Jeremy? White,?TraciHamberger,?Price?Sewel,?Bill?More,?Tim?Reed,?Phillip?Miller,?Jim?Bennet,?Jerry?Stevens,?Tim?Slone,?Joel?Beverly,?Amy?Covert,?John?Hunt,?Gareth?Turner,?Charles? Kilgore,?Michelle?Gilley,?and?Kerri?White.?Additional?thanks?are?extended?to?Troy?Best,?Gary?Hepp,?and?Robert?Lishak,?for?their?eforts?to?improve?this?research.? Funding?was?provided?by?the?United?States?Forest?Service,?Southern?Research?Station,?and?Auburn?University??additional?support?in?the?form?of?radiotransmiters?or? building?materials?was?provided?by?East?Kentucky?Power?Cooperative,?Kentucky vii? Department?of?Fish?and?Wildlife?Resources,?and?the?Daniel?Bone?National?Forest.?This?work?was?conducted?as?part?of?a?graduatedegree?at?Auburn?University. viii? Style?manual?or?journal?used:??Journal?of?Mammalogy?Computer?software?used:??Arcview?version?3.2,?Microsoft?Excel?2002,?Microsoft? Powerpoint?2002,?Microsoft?Word?2002,?and?SPSS?version?10.0 ix? TABLE?OF?CONTENTS?LIST?OF?FIGURES???????????????????????????ix? LIST?OF?TABLES??????????????????????????..xiii?CHAPTER?I:?A?REVIEW?OF?THE?CONSERVATION?AND?LIFE?HISTORY?OF? INDIANA?BATS.......................................................1?CHAPTER?I:??USE?OF?HABITAT?BY?MALE?INDIANA?BATS?..................6? ABSTRACT.......................................................11?INTRODUCTION...................................................12? MATERIALS?AND?METHODS........................................16?RESULTS.........................................................22? DISCUSSION......................................................31?LITERATURE?CITED...............................................37? CHAPTER?II:??FIDELITY?OF?MALE?INDIANA?BATS?(MYOTIS?SODALIS)?FOLLOWING?A?LARGE?SCALE?NATURAL?DISTRUBANCE.................41? ABSTRACT.......................................................41?INTRODUCTION...................................................42? MATERIALS?AND?METHODS........................................47?RESULTS.........................................................50? DISCUSSION......................................................58 x? LITERATURE?CITED...............................................63?APPENDICES......................................................65 xi? LIST?OF?FIGURES?Fig.?1.?Diagram?ofbat?boxes?constructedof?rough?cut?tulip?poplar?(Liriodendron? tulipifera)?mounted?to?3?cm?or?6?cm?diameter?rigid?conduit..???????..19?Fig.?2.?Efects?of?abundance?of?pines?and??other??trees?on?predicted?probability?of? locations?being?used?by?male?Indiana?bats?(Myotis?sodalis),?Pulaski?and?McCrearycounties,?Kentucky,?2001?2003...............................28? Fig.?3.?Efect?of?availability?of?dead?non?pine?trees?on?predicted?probability?of?locations?being?used?by?male?Indiana?bats?(Myotis?sodalis),?Pulaski?and? McCrearycounties,?Kentucky,?2001?2003...............................29?Fig.?4.?Efect?of?abundance?of?living?non?pine?trees?on?predicted?probability?of? locations?being?used?by?male?Indiana?bats?(Myotis?sodalis),?Pulaski?and?McCrearycounties,?Kentucky,?2001?2003...............................30? Fig.?5.?Declining?numbers?of?Indiana?bats?(Myotis?sodalis)?hibernating?at?South?Goldson?Cave,?Pulaski?Co.,?Kentucky,?1987?2003........................32? Fig.?6.?Fixed?kernel?estimates?for?a?population?of?male?Indiana?bats?(Myotis?sodalis)?at?SouthGoldson?Cave,?Pulaski?and?McCreary?counties,?Kentucky,? 2001?2003........................................................51?Fig.?7.?Frequency?that?male?Indiana?bats?(Myotis?sodalis)?switched?roosts,?Pulaski? and?McCreary?counties,?Kentucky,?2001?2003...........................52 Fig.?8.?Increasing?use?of?pines?(Pinus)?by?male?Indiana?bats?(Myotis?sodalis),?Pulaskiand?McCreary?counties,?Kentucky,?1996?2003.....................54? Fig.?9.?Fidelity?(number?of?days?a?bat?was?accounted?for/number?ofmovements?that?were?observed)?of?male?Indiana?bats?(Myotis?sodalis)?folowing?an?infestation? bypine?beetles?(Dendroctonus?frontalis),?Pulaski?and?McCreary?counties,?Kentucky,?2001?2003...............................................55? Fig.?10.?Fidelity?(number?of?days?a?bat?was?located/number?of?roosts?located)?of?male?Indiana?bats?(Myotis?sodalis)?to?roosts,?Pulaski?and?McCreary?counties,? Kentucky,?1996?2003...............................................56?Fig.?11.?A?conceptual?illustration?of?how?suitability?of?roost?trees?varies?over?time?? beginning?with?low?suitability?after?death?of?the?tree?(tight?bark),?folowed?by?near?maximum?suitability?(abundant?exfoliating?bark),?and?eventual?decay? (center)..........................................................57?Fig.?12.?Predicted?probability?of?roosts?remaining?suitable?for?use?by?Indiana?bats? (Myotis?sodalis)?over?time,?Pulaski?and?McCreary?counties,?Kentucky........60 xiii? LISTS?OF?TABLES?Table?1.?Number?of?male?Indiana?bats?(Myotis?sodalis)?captured,equipped?with? radiotransmiters,?and?successfully?located??1?bat?day,?Pulaski?and?McCreary?counties,?Kentucky,?2001?2003...............................23? Table?2.?Species?of?trees?used?as?roosts?by?Indiana?bats?(Myotis?sodalis),Pulaski?and?McCreary?counties,?Kentucky,?2001?2003...........................25? Table?3.?Variables?associated?with?locations?that?were?used?by?Indiana?bats?(Myotis?sodalis),?compared?to?unused?locations,?using?two?logistic?regression?models,? Pulaski?and?McCreary?counties,?Kentucky,?2001?2003.....................27 1? CHAPTER?I?A?REVIEW?OF?THE?CONSERVATION?AND?LIFE?HISTORY?OF? INDIANA?BATS?(MYOTIS?SODALIS)?The?Indiana?bat?(Myotis?sodalis)?was?placed?on?the?list?of?endangered?species?in? 1967?folowing?rapid?and?widespread?declines?in?populations?acros?most?of?its?range?(Clawson?2002).?In?1960,?the?total?population?of?Indiana?bats?was?about?883,000?? however,?the?population?had?declined?to?about?380,000?by?2001?(Clawson?2002).?Original?declines?in?populations?were?believed?to?have?been?caused?by?disturbances?in? hibernacula?by?humans?(United?States?Fish?and?Wildlife?Service?1999).?However,?populations?have?continued?to?decline?despite?construction?of?gates?at?most?large? hibernacula?(Clawson?2002).?Furthermore,?populations?of?bats?in?northern?and?southern?hibernacula?have?shown?diferent?rates?of?decline?(Clawson?2002).?These?observations? suggest?that?additional?factors?may?be?influencing?current?declines?in?populations,?including?los?of?habitats?used?during?the?non?hibernation?period,?mortality?due?to? pesticides,?or?consequences?associated?with?small?populations.?Unfortunately,?most?of?these?isues?have?not?been?the?subject?of?intensive?investigation,?with?the?posible? exception?of?habitat?used?during?the?non?hibernation?period.?Numerous?studies?have?investigated?sumer?habitat?use?byIndiana?bats? throughout?their?range?(Calahan?1993,?Calahan?et?al.?1997,?Gardner?et?al.?1991,?Gumbert?2001,?Humphrey?et?al.?1977,?Kurta?et?al.?1996),?although?most?weredescriptive 2? in?nature.?Indiana?bats?occur?throughout?much?of?the?eastern?United?States?(Gardner?and?Cook?2002),?but?evidence?of?reproduction?has?occurred?mainly?in?the?Midwest?(Clawson? 2002),?i.e.,?Illinois,?Indiana,?western?Kentucky,?southern?Michigan,?and?Misouri?(Gardner?and?Cok?2002).?This?region?frequently?has?ben?referred?to?as?the?core?range.? Habitat?in?the?region?is?often?a?mosaic?of?agricultural?lands?bordered?by?remnants?of?bottomland?forest.?Maternity?colonies?typically?occur?in?patches?of?bottomland?forests?or? fencerows,?and?individuals?usuallyare?captured?while?foraging?or?traveling?along?edges?of?agricultural?fields?(Gardner?et?al.?1991,?Kurta?et?al.?2002).?Although?now?dominated? by?extensive?agricultural?crops,?much?ofthe?region?would?have?been?prairie?before?1800?(Gardner?and?Cok?2002).? The?importance?of?bottomland?forests?to?female?and?young?Indiana?bats,?combined?with?widespread?destruction?of?wetlands,?suggest?that?significant?los?of?habitat? occurred?before?declines?in?populations?were?recognized?during?the?1960s.?Also,?large?amounts?of?pesticides?used?throughout?the?core?range?of?Indiana?bats?makes? bioaccumulation?of?contaminants?a?potential?threat.?Geluso?et?al.?(1976)?and?Clark?and?Rattner?(1987)?documented?mortality?in?bats?associated?with?specific?contaminants.? Additional?sublethal?efects?have?occurred?in?shrews?(Braham?and?Neal?1974)?and?bats?(Clark?and?Stafford?1981,?Swanepoel?et?al.?1999),?which?are?likely?to?decrease?survival? through?winter?(O?Shea?and?Clark?2002).?Detectable?levels?of?insecticides?and?other?contaminants?have?been?reported?from?bats?in?Misouri?(Schmidt?et?al.?2002),?including? evidence?of?mortality?of?Indiana?bats?(during?1975?1978)?due?to?insecticides?(O?Shea?and?Clark?2002).?Unfortunately,?efects?of?contaminants?on?populations?of?Indiana?bats?have? not?been?studied?adequately.?Clearly,?greater?investigation?of?various?aspects?of?ecology, 3? including?habitat?requirements?and?efects?of?contaminants,?is?needed?throughout?the?range?of?Indiana?bats.? Most?studies?ofecologyof?Indiana?bats?have?focused?on?female?and?young?bats?during?sumer?(Callahan?et?al.?1997,?Carter?2003,?Humphrey?et?al.?1977,?Kurta?et?al.? 2002).?Relatively?litle?is?known?about?habits?of?male?Indiana?bats.?Males?congregate?at?hibernacula?prior?to?arrival?of?females?from?summer?maternitysites,?a?behavior?known?as? swarming?(Barbour?and?Davis?1969,?Schowalter?1980).?Several?activities?occur?during?the?swarming?period,?including?mating?and?accumulation?of?fat?reserves?for?winter? (Schowalter?1980,?Thomson?1982).?Hibernation?occurs?in?>300?caves?scatered?throughout?the?eastern?United?States??however,?colonies?of>30,000?bats?occur?at?7? priority?1?hibernacula,?in?Indiana,?Kentucky,?and?Misouri?(Clawson?2002).?Bats?wintering?in?priority?1?hibernacula?represent?ca.?52%?of?the?population?(Clawson?2002).? Hibernation?occurs?in?caves?and?abandoned?mines?with?a?narrow?range?of?temperature?and?humidity?(Tuttle?and?Kennedy?2002).? Folowing?hibernation,?bats?emerge?in?spring?(Barbour?and?Davis?1969)?and?sexes?begin?roosting?separately?(Humphrey?et?al.?1977,?Kurta?et?al.?1996).?Female?Indiana?bats? usually?migrate?to?areas?away?from?hibernacula?to?form?aternity?colonies?(Barbour?and?Davis?1969,?Kurta?and?Murray?2002,?Mumford?and?Whitaker?1982).?Maternity?colonies? of<384?bats?have?been?documented?(Gardner?et?al.1991,?Humphrey?et?al.1977,?Thomson?1982,?M.?Watson?in?lit.).?Most?males?appear?to?roost?in?areas?surrounding?the? hibernacula?during?spring,?sumer,?and?autumn??althougha?smal?proportion?may?leave?and?roost?in?unknown?locations?(Gumbert?et?al.?2002).?Males?typically?rost?alone?or?in? smal?temporary?groups?of<3?bats?(Gumbert?2001).?Trees?used?by?male?Indiana?bats 4? tend?to?have?smaller?diameters?at?breast?height?than?trees?used?by?maternity?colonies?(6.4??86.6?cm??Gumbert?2001,?J.?R.?MacGregor?in?lit.,?versus?18?108?cm??Callahan?et? al.1997,?Kurta?et?al.1992,?1996,?M.?Watson?in?lit.).?From?one?site?in?Kentucky,?Gumbert?(2001)?and?Gumbert?et?al.?(2002)?provided?the?most?complete?description?of?rosting? behavior?of?male?Indiana?bats?to?date.?Roosts?of?both?sexes?usually?occur?under?exfoliating?bark?or?in?cavities?of?dead? or?damaged?trees?(Brady?et?al.?1983,?Gardner?et?al.?1991,?Gumbert?2001,?Humphrey?et?al.?1977,?Kurta?et?al.?1992),?as?well?as?under?bark?of?living?trees?(e.g.,?shagbark?hickory,? Carya?ovata???Gardner?et?al.?1991,?Gumbert?2001,?Humphrey?et?al.?1977).?Use?of?many?species?of?trees?has?been?documented?(Calahan?1993,?Gardner?et?al.?1991,?Gumbert?et?al.? 2002,?Kurta?and?Whitaker?1998,?Kurta?et?al.?1996)?and?selection?of?roosts?seems?to?vary?regionally.?Although?rare,?Indiana?bats?have?been?reportedto?roost?in?manmade? structures,?including?buildings?(Butchkoski?and?Hasinger?2002),?utility?poles?(V.?Brack,?pers.?com.),?bridges?(Barbour?and?Davis?1969),?and?bat?boxes?(Carter?2002).?Important? characteristics?include?temperature?regime?and?protection?from?weather?(R.?Rom??et?al.?in?lit.).?Other?characteristics?such?as?proximity?to?alternate?roosts?also?maybe?important? (Humphrey?et?al.?1977).?Like?numerous?species?of?bats?that?roost?in?dead?or?damaged?trees,?Indiana?bats?use?an?unknown?number?of?trees?and?switch?roosts?everyfew?days? (Gumbert?et?al.?2002,?Kurta?et?al.?2002).?Unfortunately,?many?aspects?of?the?ecology?of?Indiana?bats?remain?unknown.?For? example,?how?is?gene?flow?among?hibernacula?achieved??In?what?portion?of?the?populationdoes?most?mortality?occur??What?is?optimal?habitat??How?do?Indiana?bats? respond?to?natural?and?anthropogenic?disturbances??Do?individuals?maintain?social 5? bonds??Why?do?bats?switch?roosts??Until?these?kinds?of?isues?are?addressed,?declines?in?populations?of?Indiana?bats?are?likely?to?continue. 6? LITERATURE?CITED?Barbour,?R.?W.,?and?W.?H.?Davis.?1969.?Bats?of?America.?The?University?Press?of? Kentucky,?Lexington,?286?pp.?Brady,?J.?T.,?R.?K.?La?Val,?T.?H.?Kunz,?M.?D.?Tuttle,?D.?E.?Wilson,?and?R.?L.?Clawson.? 1983.?Recovery?plan?for?the?Indiana?bat.?United?States?Fish?and?Wildlife?Service,?Washington,?D.C.,?94?pp.? Braham,?H.?W.,?and?C.?M.?Neal.?1974.?The?efects?of?DDT?on?energetics?of?the?short??tailed?shrew,?Blarina?brevicauda.?Bulletin?of?Environmental?Toxicology?and? Contamination?12:32?37.?Butchkoski,?C.?M.,?and?J.?D.?Hassinger.?2002.?Ecology?of?a?maternity?colony?roosting?in? a?building.?Pages?130?142?inThe?Indiana?bat:?biology?and?management?of?an?endangered?species?(A.?Kurta?and?J.?Kennedy,?eds.).?Bat?Conservation? International,?Austin,?Texas,?253?pp.?Calahan,?E.?1993.?Indiana?bat?summer?habitat?requirements.?M.S.?thesis,?University?of? Misouri,?Columbia,?74?pp.?Calahan,?E.,?R.?Drobney,?and?R.?Clawson.?1997.?Selection?of?summer?roosting?sites?by? Indiana?bats?(Myotis?sodalis)?in?Misouri.?Journal?of?Mammalogy?78:818?825.?Carter,?T.?C.?2002.?Bat?houses?for?conservation?of?endangered?Indiana?Myotis.?The?Bat? House?Researcher?10:1?3. 7? Carter,?T.?C.?2003.?Summer?habitat?use?of?roost?trees?by?the?endangered?Indiana?bat?(Myotis?sodalis)?in?the?Shawnee?National?Forest?of?southern?Ilinois.?Ph.D.? disertation,?Southern?Illinois?University,?Carbondale,?82?pp.?Clark,?D.?R.,?Jr.,?and?B.?A.?Rattner.?1987.?Orthene?toxicity?to?litle?brown?bats?(Myotis? lucifugus):?acetylholinesterase?inhibition,?coordination?los,?and?mortality.?Environmental?Toxicology?and?Chemistry?6:705?708.? Clark,?D.?R.,?Jr.,?and?C.?J.?Stafford.?1981.?Efects?of?DDE?and?PCB?on?experimentally?poisoned?female?litle?brown?bats?(Myotis?lucifugus):?lethal?brain?concentrations.? Journal?of?Toxicology?and?Environmental?Health?7:925?934.?Clawson,?R.?L.?2002.?Trends?in?population?size?and?current?status.?Pages?2?8inThe? Indiana?bat:?biology?and?management?of?an?endangered?species?(A.?Kurta?and?J.?Kennedy,?eds.).?Bat?Conservation?International,?Austin,?Texas,?253?pp.? Gardner,?J.?E.,?J.?D.?Garner,?and?J.?E.?Hofmann.?1991.?Summary?ofMyotis?sodalis?sumer?habitat?studies?in?Illinois:?with?recomendations?for?impact?assessment.? Ilinois?Natural?History?Survey,?Illinois?Department?of?Conservation,?Champaign,?28?pp.? Gardner,?J.?E.,?and?E.?A.?Cook.?2002.?Seasonal?and?geographic?distribution?of?potential?sumer?habitat.?Pages?9?20inThe?Indiana?bat:?biology?and?management?of?an? endangered?species?(A.?Kurta?and?J.?Kennedy,?eds.).?Bat?Conservation?International,?Austin,?Texas,?253?pp.? Geluso,?K.?N.,?J.?S.?Altenbach,?and?D.?E.?Wilson.?1976.?Bat?morality:?pesticide?poisoning?and?migratory?stress.?Science?194:184?186. 8? Gumbert,?M.?W.?2001.?Seasonal?roost?tree?use?by?Indiana?bats?in?the?Somerset?Ranger?District?of?the?Daniel?Bone?National?Forest,?Kentucky.?M.S.?thesis,?Eastern? Kentucky?University,?Richmond,?136?pp.?Gumbert,?M.?W.,?J.?M.?O?Keefe,?and?J.?R.?MacGregor.?2002.?Roost?fidelity?in? Kentucky.?Pages?143?152?inThe?Indiana?bat:?biology?and?management?of?an?endangered?species?(A.?Kurta?and?J.?Kennedy,?eds.).?Bat?Conservation? International,?Austin,?Texas,?253?pp.?Humphrey,?S.?R.,?A.?R.?Richter,?and?J.?B.?Cope.?1977.?Summer?habitat?and?ecology?of? the?endangered?Indiana?bat,?Myotis?sodalis.?Journalof?Mammalogy?58:334?346.?Kurta,?A.,?D.?King,?J.?A.?Teramino,?J.?M.?Stribley,?and?K.?J.?Williams.?1992.?Summer? roosts?of?the?endangered?Indiana?bat?(Myotis?sodalis)?on?the?northern?edge?of?its?range.?The?American?Midland?Naturalist?129:132?138.? Kurta,?A.,?and?J.?O.?Whitaker,?Jr.?1998.?Diet?of?the?endangered?Indiana?bat?(Myotis?sodalis)?on?the?northern?edge?of?its?range.??The?American?Midland?Naturalist? 140:280?86.?Kurta,?A.,?K.?J.?Williams,?and?R.?Mies.?1996.?A?maternity?rost?of?the?endangered? Indiana?bat?(Myotis?sodalis)?in?an?unshaded,?holow,?sycamore?tree?(Platanus?occidentalis).?The?American?Midland?Naturalist?130:405?407.? Kurta,?A.,and?S.?W.?Murray.?2002.?Philopatry?and?migration?of?banded?Indiana?bats?(Myotis?sodalis)?and?efects?ofradio?transmiters.?Journal?of?Mammalogy?83:585?? 589. 9? Kurta,?A.,?S.?W.?Murray,?and?D.?H.?Miler.?2002.?Roost?selection?and?movements?acrosthe?sumer?landscape.??Pages?118?129inThe?Indiana?bat:?biology?and? management?of?an?endangered?species?(A.?Kurta?and?J.?Kennedy,?eds.).?Bat?Conservation?International,?Austin,?Texas,?253?pp.? Mumford,?R.?E.,?and?J.?O.?Whitaker,?Jr.?1982.?Mammals?of?Indiana.?Indiana?University?Press,?Blomington,?537?pp.? O?Shea,?T.?J.,?and?D.?R.,?Clark,?Jr.?2002.?An?overview?of?contaminants?in?bats,?with?special?reference?to?insecticides?and?the?Indiana?bat.?Pages?237?248?inThe? Indiana?bat:?biology?and?management?of?an?endangered?species?(A.?Kurta?and?J.?Kennedy,?eds.).?Bat?Conservation?International,?Austin,?Texas,?253?pp.? Schmidt,?A.?C.,?K.?Tyrell,?and?T.?Glueck.?2002.?Environmental?contaminants?in?bats?colected?from?Misouri.?Pages?228?236?inThe?Indiana?bat:?biology?and? management?of?an?endangered?species?(A.?Kurta?and?J.?Kennedy,?eds.).?Bat?ConservationInternational,?Austin,?Texas,?253?pp.? Schowalter,?D.?B.?1980.?Swarming,?reproduction,?and?early?hibernation?ofMyotis?lucifugus?and?M.?volans?in?Alberta,?Canada.?Journal?of?Mammalogy?61:350? ?354.?Swanepoel,?R.?E.,?P.?A.?Racey,?R.?F.?Shore,?and?J.?R.?Speakman.?1999.?Energetic?efects? of?lindane?and?environmental?factors?on?pipistrele?bats?(Pipistrellus?pipistrellus).?Environmental?Polution?104:169?177. 10? Thomson,?C.?E.?1982.?Myotis?sodalis.?Mammalian?Species?163:1?5.?Tuttle,?M.?D.,?and?J.?Kennedy.?2002.?Thermal?requirements?during?hibernation.?Pages? 68?78?inThe?Indiana?bat:?biology?and?management?of?an?endangered?species?(A.?Kurta?and?J.?Kennedy,?eds.).?Bat?Conservation?International,?Austin,?Texas,? 253?pp.?United?Sates?Fish?and?Wildlife?Service.?1999.?Agency?draft:?Indiana?bat?(Myotis? sodalis)?revised?recovery?plan.?United?States?Fish?and?Wildlife?Service,?Fort?Snelling,?Minnesota,?53?pp. 11? CHAPTER?I?USE?OF?HABITAT?BY?MALE?INDIANA?BATS?(MYOTIS?SODALIS)? ABSTRACT?The?Indiana?bat?(Myotis?sodalis)?was?placed?on?the?list?of?endangered?species?in? 1967?folowing?rapid?and?widespread?declines?in?population?acros?most?of?its?range.?Most?research?has?focused?on?maternity?colonies?and?relatively?litle?is?known?about? males.?Mortality?of?pine?trees?(Pinus)?folowing?a?large?infestation?of?southern?pine?beetles?(Dendroctonus?frontalis)?provided?an?opportunity?to?gain?insight?into?ecological? requirements?ofmale?Indiana?bats?in?Kentucky.?Mystudy?described?roosts?used?by?male?Indiana?bats,?attempted?to?determine?if?size?of?artificial?roosts(bat?boxes)?influenced?use? byother?species?of?snag?roosting?bats,?and?compared?habitats?used?by?Indiana?bats?with?unused?habitats?at?the?site.??Most?roosts?were?under?exfoliating?bark?of?dead?pines?ca.?9?m? from?the?ground?and?were?positioned?insunny?or?partially?shaded?conditions.?Bat?boxes?were?used?infrequently?(7?of?46?boxes)?by?surrogate?species?(Myotis?septentrionalis?and? Eptesicus?fuscus),?primarily?during?sumer.?Indiana?bats?used?habitats?1)that?were?dominated?by?pines?with?few??other??trees?present,?and?2)?had?greater?mean?diameters?at? breast?height,?greaterbasal?area,?more?abundant?dead?hardwoods,?and?less?abundant?living?hardwoodsthan?random?locations.?Results?indicated?open,?sunny?conditions?at? roost?trees?are?important,?posiblyfor?thermoregulation.?This?may?explain?selection?of?habitats?with?mature?trees?and?open?conditions.?Abundant?dead?hardwood?trees?in?used 12? habitats?suggests?1)?male?Indiana?bats?use?areas?with?many?potential?alternate?roosts,?and?2)?hardwoodtrees?may?have?been?used?frequently?before?natural?disturbances?killed?large? numbers?of?pine?trees.?Results?add?to?a?growing?body?of?evidence?suggesting?Indiana?bats?may?be?adapted?for?savanna?like?woodlands. 13? INTRODUCTION?The?Indiana?bat?(Myotis?sodalis)?was?placed?on?the?list?of?endangered?species?in? 1967?folowing?rapid?and?widespread?declines?in?populations?acros?most?of?its?range?(Clawson?2002).?Original?declines?in?populations?were?believed?to?have?been?caused?by? disturbances?ofhibernacula?by?humans?(United?States?Fish?and?Wildlife?Service?1999).?However,?populations?have?continued?to?decline?despite?protection?of?most?large? hibernacula?from?intrusions?by?humans?(Clawson?2002).?Factorsoccurring?during?the?non?hibernation?period?may?be?contributing?to?continued?declines?in?populations?of? Indiana?bats.?Numerous?published?and?unpublished?reports?have?investigated?ecology?of? Indiana?bats,?but?most?were?descriptions?of?rosts?used?by?maternity?colonies?during?sumer?(Callahan?et?al.?1997,?Carter?2003,?Humphrey?et?al.?1977,?Kurta?et?al.?2002).? Few?studies?have?compared?used?versus?unused?habitats,?and?biology?of?male?Indiana?bats?is?less?understood?than?that?of?females?and?young.?Carter?et?al.?(2002)?and?Gardner? and?Cok?(2002)?are?the?only?published?studies?to?have?modeled?use?of?habitat?by?Indiana?bats.?The?study?by?Gardner?and?Cok?(2002)?useda?geographic?information?system? (GIS)?to?model?habitat?throughout?the?range?of?Indiana?bats??Carter?et?al.?(2002)?modeled?habitat?of?females?in?Illinois.?Results?of?both?studies?indicated?associations?with? relatively?broad?types?of?habitat?(Carter?et?al.?2002,?Gardner?and?Cook?2002).?Gumbert 14? (2001)?provided?the?only?quantitative?analysis?of?selection?of?habitat?by?male?Indiana?bats.?Work?by?Gumbert?(2001)?and?Gumbert?et?al.?(2002)?at?South?Goldson?Cave,? Pulaski?Co.,?Kentucky,?reported?male?Indiana?bats?roosted?in?core?areas?surrounding?the?hibernaculum.?Unlike?results?from?other?parts?ofthe?range?(Calahan?et?al.?1997,?Carter? 2003,?Humphrey?et?al.?1977,?Kurta?et?al.?2002),Indiana?bats?at?South?Goldson?Cave?roosted?indead?pines?(Pinus).?Roosts?primarily?occurred?on?tops?of?ridges?and?slopes?that? were?not?steep,?and?were?closer?to?the?hibernaculum?and?ponds?than?randomlocations?(Gumbert?2001).?Variables?such?as?availability?of?potential?rosts,?basal?area,?and?tree? diameter?(dbh)?were?not?assessed.?Characteristics?of?roosts?of?many?species?of?bats?appear?related?to?maintaining? suficient,?and?stable,?temperatures?(Altringham?1999).?This?may?explain?why?maternity?colonies?typically?occur?in?roosts?and?climates?that?provide?warm?and?stable? temperatures,?especially?in?temperate?regions?(Altringham?1999,?Louren?o?and?Palmeirim?2004).?For?example,?maternity?colonies?ofIndiana?bats?oftenoccur?in?trees?with?larger? diameters?than?trees?used?by?males?(6.4?86.6?cmfor?males??Gumbert?2001,?J.?R.?MacGregor?in?lit.?18?108?cmfor?maternity?colonies??Calahan?et?al.1997,?Kurta?et? al.1992,?1996,?M.?Watson?in?lit.),?which?are?assumed?to?maintain?more?stable?temperatures.? Similar?trends?have?ben?observed?using?artificial?roosts?(bat?boxes).?Results?of?studies?of?other?species?of?bats?indicate?that?temperature?and?proximity?to?other?roosts? influence?use?by?bats?(Dillinghamet?al.?2003,?Louren?o?and?Palmeirim2004).?Bat?boxes?have?been?constructed?for?several?species?of?bats?in?North?America,?with?mixed?success? (Carter?2002,Dillingham?et?al.?2003,?Neilson?and?Fenton?1994).?Indiana?bats?generally 15? do?not?use?bat?boxes??however,?at?least?one?large?colony?was?observed?using?a?rocket?box?style?of?artificial?rost?in?Illinois?(Carter?2002).?Rocket?boxes,?a?design?with?a?continuous? crevice?around?al?4?sides?that?mimics?exfoliating?bark,?are?used?frequentlyby?a?sympatric?species,?the?northern?long?eared?bat?(Myotis?septentrionalis??pers.?observ.).? In?1999?and?2000,?forests?in?Kentucky?experienced?an?infestation?of?southern?pine?beetles?(Dendroctonus?frontalis)?that?kiled?many?pine?trees.?Mortality?of?pines?around? South?Goldson?Cave?was?not?quantified,?althoughmortalitywas?known?to?be?high.?Trees?killed?during?the?infestation?were?expectedto?create?an?abundant?supply?of?roosts?for? male?Indiana?bats?around?South?Goldson?Cave.?Combined?with?information?from?previous?studies,?the?infestation?ofsouthernpine?beetles?provided?an?opportunity?to?gain? insight?into?the?ecology?of?male?Indiana?bats.?Objectives?of?this?study?were?to?identify?factorsassociated?with?habitats?ofmale? Indiana?bats.?Two?levels?of?habitat?were?assessed,?including?microhabitat?(roosts)?and?macrohabitat?(habitat?surrounding?roosts).?Microhabitat?was?investigated?using?1)?natural? roosts?of?Indiana?bats?and?2)?an?experiment?involving?rocket?boxes?and?sympatric?species?as?surrogates?of?Indiana?bats.?Macrohabitat?was?studied?by?comparing?habitat? surrounding?roost?trees?to?habitat?surrounding?locations?not?used?by?Indiana?bats. 16? MATERIALS?AND?METHODS?The?study?area?surrounds?South?Goldson?Cave,?a?hibernaculum?that?supported?ca.? 317?Indiana?bats?in?1987,?but?contained?ca.?185?Indiana?bats?during?recent,?semiannual,?estimates?of?populationsize?during?midwinter?(T.?Wethington?pers.?com.).?A?nearby? cave?(North?Goldson?Pit,?Pulaski?Co.,?Kentucky)?is?used?sporadically?as?a?night?and?day?roost?by?Indiana?bats,?although?not?during?hibernation.?During?the?non?hibernation? period,?male?Indiana?bats?roost?in?the?forest?surrounding?the?caves?(Gumbert?et?al.?2002).?The?study?area?is?in?the?Misisippian?Plateau?and?Pottsville?Escarpment? physiographic?regions?(Wharton?and?Barbour?1973).?Topography?is?characterized?by?a?relatively?dense?network?of?steep?hills?and?valleys?drained?by?tributaries?of?the? Cumberland?River.?Cliff?lines?and?rock?overhangs?are?comon?along?ridges,?as?are?caves?and?sinkholes?in?valeys.?Forests?in?the?region?are?divided?by?the?Pottsville?Escarpment,? with?western?mesophytic?forest?in?the?Misisippian?Plateau?and?mixed?mesophytic?forest?to?the?east?(Wharton?and?Barbour?1973).?Vegetation?in?valleys?typicallyis?mixed? mesophytic?forests??whereas,?ridge?tops?and?south?facing?slopes?support?xeric?forests?dominated?byoaks?(Quercus)?and?pines?(Wharton?and?Barbour?1973).?Land?surrounding? South?Goldson?Cave?and?North?Goldson?Pit?is?part?of?the?Daniel?Bone?National?Forest,?with?scatered?privately?owned?parcels?that?are?mostly?unforested?(land?belonging?to?the? United?States?Forest?Service?=?64%??Gumbert?2001).?Eforts?to?capture?Indiana?bats?were?conducted?in5?periods?during?2001?2004, 17? including?autumn?2001?(28?29?September),?sumer?2002?(30?June?27?July),?and?spring?(12?April),?summer?(8?24?June),?and?autumn?2003?(1?5?October).?Bats?were?captured? using?a?harp?trap?(with?plastic?mesh)?and?mist?nets?outside?of?South?Goldson?Cave.?A?hand?net?also?was?used?to?capture?bats?at?activity?areas?inside?the?hibernaculum?and?North? Goldson?Pit.?The?harp?trap?and?plastic?mesh?were?placed?acros?the?entrance?of?South?Goldson?Cave?and?mist?nets?(9?by?6?m)?were?erected?ca.10?m?in?front?of?the?cave.? Additional?attempts?to?capture?Indiana?bats?were?conducted?during?summer?by?erecting?mist?nets?over?ponds?and?road?ruts?on?ridge?tops?(n=?10)?in?the?vicinity?of?the? hibernaculum.?Mist?nets?were?erected?15?min?before?sunset?and?operated?until?5?h?after?sunset.? Harp?traps?were?erected?before?dark?and?operated?until10?12?Indiana?bats?were?captured?or?until?activity?of?bats?decreased?(typically?4?5?h?after?dark).?Harp?traps?and?mist?nets? were?monitored?every?5?min?and?all?captured?bats?were?weighed?(to?the?nearest?0.25?g),?sex?and?age?were?determined?(adult?or?juvenile),?and?bands?were?attached?(split?ring,? rounded?lip,?aluminum?Kentucky?Department?of?Fish?and?Wildlife?Resources?bands).?Reproductive?condition?also?was?recorded?(pregnant,?lactating,?post?lactating,?or?testes? descended).?Male?Indiana?bats?were?fited?with?a?0.54?g?radiotransmiter?(Model?LB?2,? Holohil?Systems?Ltd.,?Carp,?Ontario,?Canada)?using?surgical?adhesive?(Skin?bond?Cement,?Smith?and?Nephew,?Inc.,?Memphis,?Tennesee),?then?released?at?site?of?capture.? Whenselecting?bats?for?attachment?ofradiotransmiters,?priority?was?given?to?Indiana?bats?that?had?ben?tracked?previously?by?Gumbert?et?al.?(2002)?or?during?the?present? study.?Bats?were?tracked?to?their?roost?trees?each?day?using?a?3?element?Yagi?antenna?and 18? radioreceiver?(Model?TRX?1000,?Wildlife?Materials,?Carbondale,?Illinois)?until?transmiters?were?shed?(i.e.,?10?17?days).? Roosttrees?were?plotted?using?handheld?GPS?units?(GlobalMap?100,?Lowrance?Electronics?Inc.,?Tulsa,?Oklahoma)?and?7.5?minute?topographic?maps,?then?marked?with? numbered?aluminum?tags,?and?blue?rings?were?painted?around?trunks.?Trees?were?identified?to?species?using?bark?characteristics?(when?posible),?and?clasified?as?living,? living?damaged,?or?dead.?New?roost?trees?were?inspected?to?determine?most?likely?cause?of?death?(southernpine?beetle?versus?other).?Trees?with?evidence?of?damage?bysouthern? pine?beetles?(pitch?tubes?or?S?shaped?tunnels?and?pupal?chambers?under?the?bark??United?States?Department?of?Agriculture?1981)?were?assumed?to?have?been?killed?during?the? infestation??trees?without?evidence?of?damage?bysouthernpine?beetles?were?classified?as??other.???Additionally,?diameter?at?breast?height?(dbh)?was?measured?using?a?diameter? tape?(to?the?nearest?0.1?cm)?and?height?of?tree?was?visually?estimated?(to?the?nearest?meter).?Height?of?bats?relative?to?the?ground?was?estimated?using?a?Yagi?antenna?while? standing?7?10?m?from?the?tree.?Telemetric?estimates?of?height?were?confirmed?by?visual?identification?of?bats?using?a?miror?and?binoculars?whenever?posible.?Solar?exposure?at? height?of?each?roost?was?estimated?visually?(shaded,?intermediate,?or?sunny)?because?canopy?coverage?at?ground?level?did?not?accurately?reflect?this?characteristic?(Gumbert? 2001).?Bat?boxes?(n?=?46)?were?constructed?of?rough?cut,?tulip?poplar?(Liriodendron? tulipifera),?using?a?design?modified?from?J.?R.?MacGregor?and?D.?Dourson?(in?lit.??Fig.?1).?Woodenportions?ofroosts?were?mounted?to?3?m?long?metal?poles?(3?cm 19? Fig?1.?Diagram?ofbat?boxes?constructedof?rough?cut?tulip?poplar?(Liriodendron?tulipifera)?mounted?to?3?cm?or6?cm?diameter?rigid?conduit.?Roosts?were?modified?from?a? design?by?D.?Dourson?and?J.?R.?MacGregor?(in?lit.). 20? or?6?cmdiameter?rigid?conduit)?and?anchored?in?the?ground?with?27?kg?of?concrete.?Centers?of?roosts?were?constructed?using?large?(15?by?15?cm)?or?small?(10?by?10?cm)?1?m?? long?wooden?posts.?Walls?were?constructed?from?4?cm?thick?by?83?cm?long?wooden?planks?that?were?mounted?to?center?posts,?leaving?a?2?cm?wide?cavity?around?center?posts? for?bats?to?roost.?Pitched?wooden?roofs?(22.5??angle??4?cm?thick)?were?attached?to?the?tops?of?each?box?to?exclude?precipitation.? Large?(n?=?23)?and?small?(n?=?23)?boxes?were?paired?2?m?apart?with?entrances?3.3?m?from?the?ground.?Pairs?of?boxes?were?arranged?into?groups?with?each?pair?spaced?15?m? apart.?Boxes?were?placed?at?Double?Tarkiln?Ridge?(<1?km?from?South?Goldson?Cave)?and?Goodwater?(a?reclaimed?strip?mine?ca.?64?km?northeast?of?South?Goldson?Cave,? Pulaski?Co.,?Kentucky)?from?August?2002throughOctober?2004.?Surveys?of?boxes?were?conducted?during?April,?June,?and?October?2003,?and?March?and?October?2004?to? document?use?by?bats.?Use?of?artificial?rosts?by?bats?was?determined?by?presence?of?guano?at?bases?of?roosts?and?by?visually?inspecting?cavities?for?bats?(using?a?small?miror? to?reflect?sunlight?into?cavities).?When?posible,?numbers?and?species?of?bats?were?recorded,?or?amounts?of?guano?were?estimated?(low,?moderate,?or?high).? Data?describing?macrohabitat?characteristics?were?colected?from?the?area?surrounding?64?roost?trees?using?a?10?factor?prism?(Avery?1967),?while?standing?1?m? from?the?roost?(Gumbert?2001).?Nearby?trees?that?were?not?fully?displaced?using?the?prism?were?counted,?identified?to?species,?clasified?as?living?or?dead,?measured?using?a? diameter?tape,?and?basal?area?of?the?habitat?surrounding?each?roost?was?calculated?(Avery?1967).?Dead?pines?were?classified?according?to?cause?of?death?(pine?beetle?or?other).?The? same?methods?were?used?to?describe?the?study?area,?using?10?randomly?placed?transects 21? (ca.?200?250?m?long),?during?July?2002.?Transects?were?placed?within?a?5?km?radius?of?the?hibernaculum?and?each?contained?6?8?evenly?spaced?non?overlapping?points?where? data?were?colected?(n=?63).?Transects?occurred?outside?of?core?areas?identified?by?Gumbert?et?al.?(2002).? Logistic?regression?models?were?used?to?identify?characteristics?of?macrohabitat?that?explained?significant?amounts?of?variation?in?use?of?habitat?by?Indiana?bats.?Related? variables?were?separated?into?2?suites?of?variables,?including?abundance?of?trees?of?diferent?genera?(numbers?of?pines,?oaks,?hickories,?maples?[Acer],?or??other?),?and? characteristics?of?nearby?trees?(basal?area,?numbers?of?dead?pines,?living?pines,?dead?non??pines,?and?living?non?pines).?A?separate?logistic?regression?model?was?used?to?analyze? each?suite?of?variables.?Significant?variables?were?identified?using?the?backwards??elimination?procedure,?which?selected?variables?that?improved?the?predictive?accuracy?of? logistic?regression?models?and?excluded?those?that?did?not?("of?removal?=?0.05).?Analyses?were?conducted?using?SPSS?10.0?at "=?0.05?(Green?et?al.?2000). 22? RESULTS?Bats?captured?and?banded?at?the?entrance?and?inside?of?South?Goldson?Cave?and? North?Goldson?Pit?included?the?Rafinesque?s?big?eared?bat?(Corynorhinus?rafinesquii?n?=?11),?big?brown?bat?(n?=?7),?eastern?red?bat?(Lasiurus?borealis?n?=?1),litle?brown?bat?(n? =?117),?northern?long?eared?bat?(n=?144),?Indiana?bat?(n?=?48),?evening?bat?(Nycticeus?humeralis?n?=?1),?and?eastern?pipistrele?(Pipistrellus?subflavus?n?=?155).?In?general,?the? same?species?of?bats?captured?at?caves?also?were?capturedover?ponds,?except?no?litle?brown?bats?or?Indiana?bats?were?caught,?and?relatively?few?eastern?pipistreles?were? caught.?Bats?captured?over?ponds?included?the?Rafinesque?s?big?eared?bat?(n=?1),?big?brown?bat?(n?=?13),?eastern?red?bat?(n=?7),?northern?long?eared?bat?(n=?91),?evening?bat? (n?=?1),?and?eastern?pipistrele?(n?=?4).?Male?Indiana?bats?were?captured?(n=?34),?equipped?with?radiotransmiters?(n=? 31),?and?located?(n=?25)?during?4?sampling?periods?(Table?1).?Female?Indiana?bats?(n?=15)?were?captured?using?a?hand?net?during?spring?and?autumn?2003,?but?were?not? equipped?with?radiotransmiters?because?females?typically?enter?hibernation?earlier?than?males?(Brady?et?al.?1983,?J.?R.?MacGregor?in?lit.)?or?migrate?from?the?study?area? folowing?hibernation?(Gumbert?2001).?Succes?in?capturing?bats?varied?seasonally,?with?relatively?low?succes?during?sumer?(Table?1). 23? Table?1.?Number?ofmale?Indiana?bats?(Myotis?sodalis)?captured,?equipped?with?radiotransmiters,?and?successfully?located??1?bat?day?a?,?Pulaski?and?McCreary?counties,? Kentucky,?2001?2003.?Tracking?period? Captures? With?radiotransmiters? Located? Autumn?2001?10? 9? 7?Sumer?2002?5? 5? 5? Spring?2003?b? 11? 9? 7?Sumer?2003?0? 0? 0? Autumn?2003?9? 9? 7?Total? 34? 31?c? 25?c? a?A?bat?day?represents?1?bat?located?during?1?day.?b?Radiotransmiters?were?not?affixed?to?2?bats?because?of?low?body?weights? (radiotransmiters?weighed?>9%?ofbody?weight).?c?Included?a?bat?that?was?radiotrackedtwice?(autumn?2001?and?2003). 24? Male?bats?were?observed?using?87?roost?trees?on?256?occasions.?Pines?were?the?predominant?trees?used?by?bats,?but?relative?proportion?of?trees?appeared?to?vary? seasonally,?with?lowest?use?of?pines?occurring?in?spring?2003?(Table?2).?Bats?roosted?alone?or?in?small?groups?(2?3?bats),?usually?in?dead?trees?(92%?of?roosts?were?dead?? Appendix?1).?Most?(n=?74?of?87)?roost?sites?wereunder?exfoliating?bark.?Other?roost?sites?were?in?boles?of?trees?that?were?splintered?at?the?top?(n=?3),?crevices?(n?=?1),?or? unknown?locations?(n?=?5).?Average?height?of?roost?sites?was?9???0.5?m(range?=?3?21?m??Appendix?1).?Individuals?that?roosted?lower?to?the?ground?were?easier?to?confirm?visually? than?those?higher?up.?Seventeenroosts?were?confirmed,?but?no?bat?was?seen?>12?m?from?the?ground.?Locations?where?bats?roosted?on?trees?usually?had?sunny(n?=?49?of?81? roosts)or?intermediate?(n?=?22)exposure?to?sunlight??relatively?few?(n?=?10)?were?shaded.?Northern?long?eared?bats?were?the?only?species?observed?using?artificial?rosts?? however,?some?large?sized?guano?underneath?roosts?suggested?use?by?big?brown?bats.?Use?of?artificial?rosts?by?bats?was?limited?to?1?large?roost?(of?21?roosts?available)?in? April?2003,?and?5?smal?and?1?large?roost?(of?21?roosts?available)?during?June?2003.?Solitary?bats?were?observed?roosting?in?3?roosts?and?moderate?amounts?of?guano?were? observed?under?the?remaining?4.?No?additional?evidence?of?use?was?observed?during?subsequent?surveys.?Statistical?analysis?was?not?performed?due?to?the?large?number?of? roosts?(n=?39?of?46)?that?were?not?used?by?bats.?Data?colected?at?points?along?random?transects?in?sumer?2002?indicated?the? forest?surrounding?South?Goldson?Cave?was?33??3%oaks,?18??3%pines,?16???2%?maples,?11???2%hickories,?and?23???3%other?(Appendix?2).?Twentypercent?(106of? 539)?trees?surveyed?were?dead.?Pines?were?the?most?abundant?species?of?dead?trees 25? Table?2.?Species?of?trees?used?as?roosts?by?male?Indiana?bats?(Myotis?sodalis),?Pulaski?and?McCreary?counties,?Kentucky,?2001?2003.?Usage?is?expresed?as?number?of?trees? used?and?percentage?of?the?total?number?of?trees?used?(in?parentheses)?within?each?radiotelemetric?monitoring?period.? Taxa? Autumn?2001?Sumer?2002?Spring?2003?Autumn?2003?Total?Acer?rubrum? 1?(3%)? 1?(5%)? 2?(3%)? Carya?ovata? 1?(3%)? 2?(10%)? 3?(5%)?Pinus? 20?(69%)? 9?(100%)? 14?(67%)? 27?(96%)? 70?(72%)? rigida? 4? 2? 6?echinata? 14? 8? 11? 24? 57?virginiana? 2? 1? 3? 1? 7? Quercus? 6?(21%)? 2?(10%)? 1?(4%)? 9?(15%)?alba? 1? 1? 1? 3?borealis? 3? 1? 4? velutina? 2? 2?Fraxinus? 1?(3%)? 1?(2%)? Unknown? 2?(10%)? 2?(3%)?Total? 29? 9? 21? 28? 87 26? (92%,?97?of?106)?compared?to?non?pines?(8%,?9?of106).?Nearly?all?pines?were?dead?(93%,?97?of?104)?however,?59%?(57?of?97)?of?those?appeared?to?have?been?killed?by? southernpine?beetles?and?41%?(40?of?97)?showed?no?evidence?of?damage?from?pine?beetles.?Overal,?55%?(57?of?104)?of?al?pines?appeared?to?have?been?killed?by?pine? beetles.?Logistic?regressionanalysis?indicated?that?roost?trees?occurred?in?macrohabitats? (Appendix?3)?that?difered?fromrandomlocations?in?the?study?area?(Table?3).?Specifically,?analysis?using?abundance?of5?categories?oftree?species?(model?1??Table?3)? indicated?bats?roosted?in?habitats?that?were?dominated?by?pines?and?had?fewer??other??trees?present?(Fig.?2).??Other??trees?included?numerous?species,?many?of?which?were? comon?in?the?understory?or?were?not?associated?with?xeric?ridges?(e.g.,Liriodendron?tulipifera,?Liquidambar?styraciflua,?Prunus).?Used?and?available?locations?were? clasified?corectly?68%?of?the?time?using?this?model.?Other?macrohabitat?variables?also?were?associated?with?used?locations,?including?mean?dbh,?basal?area,?numbers?of?dead? non?pines,?and?numbers?of?living?non?pines?(model?2?Table?3).?Bats?roosted?in?habitats?with?greater?numbers?of?dead?non?pines?(Fig.?3),?mean?dbh,?and?basal?area,?than?unused? locations.?Also,?used?habitats?were?negativelyasociated?with?living?non?pines?(Fig.?4).?Locations?were?classified?corectly?81%?of?the?time?using?this?model. 27? Table?3.?Variables?associated?with?locations?that?were?used?by?Indiana?bats?(Myotis?sodalis),?compared?to?unused?locations,?using?two?logistic?regression?models,?Pulaski?and? McCreary?counties,?Kentucky,?2001?2003?a?.?Model? Variable?b? ?? W?c? Log?odds? P? 1? Pine? 0.33?14.06?1.39?1?occasion.?Several?bats?(n?=?5)?received?the?added?disturbance?of? being?equipped?with?radiotransmiters?during?>1?radiotelemetry?period?prior?to?this?study,?including?1?bat?that?was?tracked?5?times?from?October?1998?through?September?2002? (Gumbert?2001,?M.?W.?Gumbert?pers.?com.,?Gumbert?et?al.?2002).?During?the?present?study,?this?particular?bat?was?recaptured?in?a?mist?net?and?equipped?with?a? radiotransmiter?on?2?October?2003,?and?recaptured?at?South?Goldson?Cave?3?nights?later.?To?my?knowledge,?this?bat?has?ben?recaptured?(n=?6?times)?and?located?(n=?6?periods)? more?times?than?any?other?Indiana?bat.?These?data?support?results?by?Kurta?and?Murray?(2002),?who?reported?relatively?high?rates?of?recapture.? Characteristics?of?roost?trees?were?similar?to?those?reported?by?Gumbert?(2001),?where?bats?roosted?in?pine?treesthat?had?full?or?intermediate?exposure?to?sunlight.? Although?roosts?were?in?uplands?and?most?occurred?in?pines,?versus?hardwood?trees?in 34? bottomlands,?results?supported?trends?in?other?regions?where?roosts?typically?occur?in?sunny?microhabitats?(Calahan?1993,?Calahan?et?al.?1997,?Gardner?et?al.?1991,?Humphrey? et?al.?1977,?Kurta?et?al.?1996).?This?supports?the?hypothesis?that?warm?microclimates?are?important?to?many?species?of?bats?in?temperate?regions?(Altringham?1999,?Louren?o?and? Palmeirim2004),?including?Indiana?bats?(Humphrey?et?al.?1977),?and?may?explain?why?artificial?roosts?were?occupied?mostlyduring?summer.? Selection?of?habitat?by?male?Indiana?bats?provides?more?evidence?to?suggest?the?importance?of?temperature?regime?to?roosting?bats.?Bats?may?have?roosted?in?stands?that? were?dominated?by?pines?and?had?few?species?oftrees?comon?in?the?understorybecause?of?open,?sunny?conditions?in?those?stands.?In?Kentucky,?habitats?dominated?by?pines? typically?occur?on?xeric?ridges?(Wharton?and?Barbour?1973),?where?Gumbert?(2001)?determined?male?Indiana?bats?were?most?likely?to?occur.?Selection?ofhabitats?with? greateraverage?dbh,?greaterbasal?area,?and?few?living?hardwood?trees,?may?have?contributedto?warm?and?stable?temperatures?at?roosts.? Where?male?Indiana?bats?forage?in?relation?to?core?areas?is?not?available?in?published?literature.?Females?and?juveniles?may?forage?<1?2.5kmof?rosting?sites,?along? edges?of?woodlots,?and?stream?and?road?coridors?(Brady?et?al.?1983,?Humphrey?et?al.?1977,?Mumford?and?Cope?1958).?During?the?present?study,?1?male?bat?was?monitored? and?remained?in?the?vicinity?(ca.?30,000?hibernating?bats?United?States?Fish?and?Wildlife?Service?1999).?However,?Indiana?bats?hibernate?in?several?smaller?colonies?throughout?the?state?(B.?Palmer?Bal,?Jr.? pers.?com.).?Fidelity?of?male?Indiana?bats?has?been?studied?at?a?small?hibernaculum,?South?Goldson?Cave,?Pulaski?Co.,?Kentucky,?since?1996?(Gumbert?et?al.?2002).? Land?surrounding?the?cave?is?located?in?Pulaski?and?McCreary?counties,?Kentucky,?in?the?Somerset?Ranger?District?of?the?Daniel?Bone?National?Forest.? Research?conducted?by?J.?R.?MacGregor?(in?lit.)?and?Gumbert?(2001)?revealed?that?Indiana?bats?at?South?Goldson?Cave?roosted?in?several?species?of?trees,?but?pines?(Pinus)? were?used?most?often?(Gumbert?2001,?J.?R.?MacGregor?in?lit.).?Bats?switched?roosts?often,?but?most?days?were?spent?roosting?in?core?areas?scatered?around?the?hibernaculum? (Gumbert?et?al.?2002).?The?study?by?Gumbert?et?al.?(2002)?was?unique?in?that?data?were?colected?from?any?bats?(n?=?60),?and?some?bats?(n?=?16)?were?tracked?for?multiple? seasons?or?years.?Beginning?in1999?and?peaking?in2000,?the?Daniel?Boone?National?Forest? experienced?an?infestation?ofsouthernpine?beetles?that?killed?large?numbers?of?trees.?Previously,?pines?were?a?dominant?overstory?species?(Gumbert?2001).?High?mortality?of? pines?was?likely?to?influence?the?bat?comunity?in?the?Somerset?Ranger?District? 45? however,?specific?efects?could?not?be?predicted.??Increased?numbers?of?dead?pines?were?expected?to?provide?a?temporary?abundance?of?potential?roosts?around?South?Goldson? Cave.?I?hypothesized?that?pine?trees?killed?during?the?infestationwould?deteriorate?at?similar?rates?and?might?become?unsuitable?for?use?as?roosts?around?the?same?time.? Depending?on?how?quickly?rosts?deteriorated?and?availability?of?alternative?rosts,?bats?could?experience?a?shortage?of?roosts.?Unfortunately,?litle?is?known?about?dynamics?of? deterioration?and?replacement?of?standing?dead?trees?(snags)?and?the?efects?onfidelityof?bats.? Changes?in?abundance?of?dead?pines?at?the?site?were?expected?to?influence?several?species?of?bats?that?roost?indead?trees,?including??Indiana?bats,?litle?brown?bats?(Myotis? lucifugus),?northern?long?eared?bats?(Myotis?septentrionalis),?and?big?brown?bats?(Eptesicus?fuscus).?In?addition?to?influencing?distribution?of?potential?roosts,?changes?in? structure?and?dominance?of?overstory?trees?was?likely?to?influence?distribution?and?abundance?of?insect?prey.?Distribution?of?fodalso?mayinfluence?selection?of?rosts?by? bats,?although?the?trend?has?not?been?demonstrated?in?insectivorous?bats?(Lewis?1995).?Efects?of?changes?in?distribution?and?abundance?of?insect?prey?at?the?study?site?would?be? dificult?to?detect?and?predict.?Identification?of?digested?insects?is?limited?to?course?taxonomic?groups,?and?no?baseline?data?existed?at?South?Goldson?Cave.?Thus,?the? present?study?focused?on?aspects?of?fidelity?directly?related?to?abundance?and?distribution?ofroosts.? Major?objectives?of?the?current?study?were?to?determine?if?bats?used?trees?killed?during?the?infestation?ofsouthernpine?beetles,?and?to?determine?if?use?of?core?areas?and? fidelity?to?individual?rosts?were?influenced?by?increased?abundance?of?potential?rosts. 46? Specifically,?I?hypothesized?that?increased?availability?of?potential?roosts?folowing?the?infestation?would?result?in?1)?increased?use?of?pine?trees,?2)?changes?in?locations?of?core? areas,?and?3)?decreased?fidelity?to?individual?roosts.?Another?goal?was?to?measure?rates?of?deterioration?of?roosts?to?estimate?how?long?trees?may?remain?available?to?bats. 47? MATERIALS?AND?METHODS?Bats?were?captured?using?harp?traps?and?mist?nets,and?tracked?via?radiotelemetry? (Chapter?I)?using?the?methods?of?studies?conducted?prior?to?the?infestation?(Gumbert?2001,?Gumbert?et?al.?2002).?Roosts?were?plotted?and?marked?(Chapter?I)?and? coordinates?of?roosts?were?incorporated?into?a?geographic?information?system?(GIS).?Radio?telemetry?locations?were?analyzed?using?a?fixed?kernel(Powel?2000)?to?identify? home?range?(95%?contour)?and?core?areas?(50%?contours)using?the?animal?movement?extension?(Hooge?and?Eichenlaub?2000)?to?Arcview?3.2?(Environmental?Systems? Research?Institute,?Redlands,?California).?Kernel?estimates?are?based?onthe?geographic?distribution?of?radio?telemetry?locations?and?frequency?that?locations?are?used?(Powell? 2000).?Estimates?are?expressed?as?contours,?which?represent?the?percentage?of?maximum?probability?a?given?locationwill?occur?in?a?homerange?(Powell?2000).? Bats?sometimes?roosted?in?undetected?locations?before?moving?back?to?trees?that?alowed?radiotelemetric?detection.?When?this?happened,?bats?were?either?too?far?from?the? study?area?to?be?detected,?or?were?hidden?by?landscape?features?that?made?detection?unlikely?(e.g.,?ridges,?steep?valleys,?caves).?I?argue?that?these?movements?are?meaningful? and?should?be?considered?when?estimating?fidelity.?Thus,?fidelityestimates?for?individual?bats?were?calculated?by?dividing?number?of?days?a?bat?could?be?accounted?for?by?number? of?movements?observed?during?that?period.?In?some?cases,?recorded?movements?included?instances?when?bats?moved?from?documented?roosts?to?unknown?locations.?This 48? approach?was?used?to?describe?changes?in?fidelity?during?the?present?study?(2001?2003).?J.?R.?MacGregor?(in?lit.)?used?the?same?method?to?calculate?fidelity?during?his?work?at? South?Goldson?Cave?during?1996?1997,?but?reported?average?values.?Thus,?comparisons?between?studies?had?to?be?conducted?using?a?less?accurate?method?of?calculating?fidelity? (number?of?days?bats?were?located?divided?by?number?of?trees?located).?This?method?fails?to?take?into?account?instances?when?bats?move?to?unknown?locations,?but?alowed? combining?of?results?for?1996?1997?and?2001?2003.?Fidelity?data?from?1998?1999?(Gumbert?et?al.?2002)?were?not?reported?in?a?format?that?allowed?comparisons.? Subsamples?of?roost?trees?(n?=?59)?identified?during?this?study?orby?Gumbert?et?al.?(2002)?were?revisited?during?spring?and?autumn?2003?and?spring?and?autumn?2004?to? determine?how?long?trees?remained?suitable?for?use?by?bats.?Trees?that?were?still?standing?with?bark?or?crevices?were?considered?suitable?for?use?as?roosts??trees?on?the?ground?or? standing?with?no?obvious?bark?or?crevices?were?considered?unsuitable?for?use?as?roosts.?Time?was?measured?in?months?from?the?date?a?tree?was?first?located?until?the?tree?was? observed?as?unsuitable.??If?a?tree?remained?suitable,?time?was?measured?as?number?of?months?from?the?date?first?located?until?the?most?recent?survey.? Three?linear?regression?models?were?used?to?detect?potential?changes?in?roosting?behavior?over?time.??One?model?determined?if?Indiana?bats?used?increasing?proportions?of? pines?during?1996?2003,?using?average?values?from?J.?R.?MacGregor?(in?lit.),?Gumbert?(2001),?and?the?present?study.?The?remaining?2?models?were?used?to?determine?if?fidelity? changed?over?time.??Specifically,?potential?changes?in?fidelity?(number?ofdays?a?bat?was?observed?divided?by?number?of?movements?that?were?observed)?during?2001?2003?were? plottedusing?1?model,?and?a?separate?regression?model?was?used?to?plot?fidelity?(number 49? of?days?bats?were?located?divided?by?number?of?trees?that?were?located)?for1996?1997?and?2001?2003.?Additionally,?logistic?regressionanalysis?was?used?to?determine? probability?of?roosts?remaining?suitable?over?time.??Variables?included?suitability?of?roosts?(0?=?unsuitable,?1?=?suitable)?as?the?dependent?variable?and?age?(months?since? located?by?a?researcher)?and?type?of?rost?(0?=?non?pine,?1?=?pine)?as?independent?variables.?Logistic?regression?models?were?constructed?using?the?backwards?elimination? method,?with?variables?selected?based?on??of?removal=?0.05.?Predicted?probabilities?of?trees?remaining?suitable?over?time?were?generated?from?logistic?regression?models?(Green? et?al.?2000).?Analyses?were?conducted?using?SPSS?10.0(Green?et?al.?2000)?at???=?0.05. 50? RESULTS?Average?distance?between?roost?trees?was?644?mand?maximum?distance?was?8.5? km.?Fixed?kernel?analysis?using?24?bats?located?in?a?total?of?256?days?indicated?3?core?areas?(2?discrete?core?areas??Bat?Ridge?and?Double?Tarkiln?Ridge/Bauer)?that?were? centered?<2?km?from?South?Goldson?Cave?(Fig.?6).?The?Bauer?core?area?was?used?more?frequently?by?bats?during?the?present?studythan?during?Gumbert?et?al.?(2002)??likewise,? Bethel?Church?and?Yelowjacket?were?identified?by?Gumbert?et?al.?(2002),?but?were?not?used?as?core?areas?during?the?present?study.?Roosttrees?in?Yelowjacket?(n=?20)?were? revisited?in?2003?and?only?2?living?shagbark?hickories?(Carya?ovata)?were?observed?standing.?Remaining?roost?trees?in?Yellowjacket?had?either?fallen?(n?=?8)?or?were?not? found?(n?=?10).?In?general,?bats?moved?to?new?roosts?every?2?5?days?(Fig.?7).?Some?trees?(n?=?5)? were?used?by?>1?bat?during?the?same?radiotelemetry?period?(Appendix?3).?Vocalizations?from?roosting?bats,?as?wel?as?observations?made?visually?and?using?radiotelemetry,? indicated?bats?sometimes?used?trees?simultaneously?(in?groups?of<3?bats).?Several?trees?(n?=?4)?were?used?as?roosts?during?previous?studies,?including?a?damaged?white?oak? (Quercus?alba),?which?was?used?during?autumn?1998?2000?(Gumbert?et?al.?2002)?and?reused?by?the?same?bat?during?this?study.?Likewise,?in?spring?2003,?I?located?bat? 1929KY/A00010DB?in?the?same?living?shagbark?hickory?it?used?during?spring?1998?(M.?W.?Gumbert?pers.?com.). 51? Fig.?6.?Fixed?kernel?estimates?for?a?population?of?male?Indiana?bats?(Myotis?sodalis)?at?SouthGoldson?Cave,?Pulaski?and?McCreary?counties,?Kentucky,?2001?2003.?Contours? (95%?and?75%)?are?represented?byunshaded?and?shaded?polygons,?respectively??core?areas?(50%?contours)?are?depicted?by?diagonal?hatching.?Locations?of?roost?trees?and? transects?are?indicated?by?dots?and?diamonds,?respectively. 52? 0?1?2? 3?4?5? 6?7?8? 9?10?1? 1? 2? 3? 4? 5?Fig?7.?Frequency?that?male?Indiana?bats?(Myotis?sodalis)?switched?roosts,?Pulaski?and? McCreary?counties,?Kentucky,?2001?2003.? Days?spent?between?movements? Number?of ?bats 53? Increasing?use?of?pines?occurred?during?1996?2003?(r?2?=?0.51,F=?10.4,d.f.?=?1,?10,?P1?bat,?including?trees?458,?732,?674,?and?769,?which?were?reused?from?previous?studies. 72? Appendix?2.?Characteristics?of?habitats?not?used?by?male?Indiana?bats?(Myotis?sodalis)?in?the?vicinity?of?South?Goldson?Cave,?Pulaski?and?McCreary?counties,?Kentucky,?July?2002.? Random?location?Average?dbh?(cm)?Basal?area?(m?2?/ha)? Pinus?Quercus?Acer?Carya?Other?Number?of?tres?kiled?by?southern?pine?betles? Dead?pines? Living?pines? Dead?non?pines? Living?non?pines? 1? 34.9? 14.8? 0? 10?0?0? 2? 0? 0? 0? 0? 12?2? 24.7? 11.1? 3? 3? 2?0? 1? 2? 3? 0? 0? 6? 3? 31.3? 18.5? 2? 5? 6?2? 0? 2? 2? 0? 0? 13?4? 35.9? 14.8? 3? 6? 3?0? 0? 0? 3? 0? 0? 9? 5? 32.8? 14.8? 1? 6? 5?0? 0? 0? 1? 0? 0? 11?6? 27.9? 14.8? 4? 3? 4?0? 1? 1? 1? 3? 1? 7? 7? 22.6? 13.6? 3? 4? 3?0? 1? 0? 0? 3? 0? 8?8? 42.0? 14.8? 4? 3? 3?0? 2? 0? 1? 3? 0? 8? 9? 37.9? 17.3? 3? 7? 4?0? 0? 0? 0? 3? 0? 11?10?39.6? 16.0? 0? 6? 4?1? 2? 0? 5? 0? 0? 7? 11?27.2? 12.3? 0? 0? 3?0? 7? 0? 0? 0? 1? 9?12?27.5? 7.4? 0? 0? 1?1? 4? 0? 0? 0? 0? 5? 13?26.4? 6.2? 0? 2? 0?1? 2? 0? 0? 0? 0? 3?14?25.1? 4.9? 0? 0? 1?1? 2? 0? 0? 0? 0? 4? 15?15.4? 14.8? 7? 2? 0?0? 3? 4? 7? 0? 0? 5 73? Appendix?2?continued.? Random?location?Average?dbh?(cm)?Basal?area?(m?2?/ha)? Pinus?Quercus?Acer?Carya?Other?Number?of?tres?kiled?by?southern?pine?betles? Dead?pines? Living?pines? Dead?non?pines? Living?non?pines? 16?15.9? 11.1? 5? 1? 2?0? 1? 3? 5? 0? 0? 5?17?16.5? 13.6? 7? 2? 0?1? 1? 4? 7? 0? 0? 5? 18?22.3? 9.9? 7? 0? 0?0? 1? 7? 7? 0? 0? 1?19?14.5? 8.6? 1? 3? .? 1? 2? 0? 1? 0? 0? 6? 20?16.3? 12.3? 2? 3? 0?3? 2? 0? 2? 0? 0? 8?21?23.3? 6.2? 0? 3? 0?0? 2? 0? 0? 0? 0? 5? 22?23.2? 12.3? 5? 0? 0?1? 4? 3? 5? 0? 0? 5?23?30.0? 9.9? 2? 6? 0?0? 0? 0? 2? 0? 1? 5? 24?43.9? 4.9? 0? 1? 1?0? 2? 0? 0? 0? 0? 5?25?35.4? 8.6? 0? 2? 1?1? 3? 0? 0? 0? 0? 7? 26?29.8? 9.9? 0? 1? 1?4? 2? 0? 0? 0? 0? 8?27?31.8? 9.9? 1? 4? 0?1? 2? 1? 1? 0? 0? 7? 28?24.5? 8.6? 0? 4? 3?0? 0? 0? 0? 0? 0? 7?29?33.4? 8.6? 2? 2? 2?0? 1? 2? 2? 0? 0? 5? 30?54.0? 12.3? 9? 1? 0?0? 0? 8? 9? 0? 0? 1?31?26.0? 7.4? 0? 0? 0?1? 5? 0? 0? 0? 0? 6 74? Appendix?2?continued.? Random?location?Average?dbh?(cm)?Basal?area?(m?2?/ha)? Pinus?Quercus?Acer?Carya?Other?Number?of?tres?kiled?by?southern?pine?betles? Dead?pines? Living?pines? Dead?non?pines? Living?non?pines? 32?28.7? 8.6? 3? 0? 0?0? 4? 0? 3? 0? 0? 4?33?29.2? 14.8? 1? 3? 0?0? 8? 0? 1? 0? 0? 11? 34?42.3? 7.4? 0? 4? 0?1? 1? 0? 0? 0? 0? 6?35?39.1? 7.4? 0? 3? 1?0? 2? 0? 0? 0? 0? 6? 36?37.4? 11.1? 2? 1? 1?1? 4? 2? 2? 0? 1? 6?37?32.7? 11.1? 0? 6? 2?0? 1? 0? 0? 0? 0? 9? 38?34.5? 7.4? 0? 3? 2?1? 0? 0? 0? 0? 0? 6?39?28.1? 8.6? 0? 2? 4?0? 1? 0? 0? 0? 0? 7? 40?32.6? 8.6? 1? 3? 0?0? 3? 0? 1? 0? 1? 5?41?25.4? 7.4? 0? 2? 3?0? 1? 0? 0? 0? 0? 6? 42?18.4? 8.6? 0? 3? 2?0? 2? 0? 0? 0? 1? 6?43?23.0? 14.8? 2? 1? 4?2? 3? 0? 2? 0? 1? 9? 44?29.0? 9.9? 3? 3? 1?0? 1? 3? 3? 0? 0? 5?45?16.8? 8.6? 2? 2? 0?1? 2? 2? 2? 0? 0? 5? 46?29.7? 6.2? 2? 2? 0?0? 1? 1? 2? 0? 0? 3?47?20.2? 14.8? 7? 1? 1?0? 3? 3? 7? 0? 0? 5 75? Appendix?2?continued.? Random?location?Average?dbh?(cm)?Basal?area?(m?2?/ha)? Pinus?Quercus?Acer?Carya?Other?Number?of?tres?kiled?by?southern?pine?betles? Dead?pines? Living?pines? Dead?non?pines? Living?non?pines? 48?22.8? 8.6? 5? 1? 0?0? 1? 5? 5? 0? 0? 2?49?23.5? 8.6? 2? 3? 0?0? 2? 2? 2? 0? 0? 5? 50?28.8? 14.8? 0? 4? 4?4? 0? 0? 0? 0? 0? 12?51?35.8? 7.4? 0? 1? 0?4? 1? 0? 0? 0? 0? 6? 52?33.3? 7.4? 0? 1? 0?5? 0? 0? 0? 0? 0? 6?53?37.8? 11.1? 0? 1? 1?4? 3? 0? 0? 0? 0? 9? 54?32.7? 8.6? 0? 1? 2?1? 3? 0? 0? 0? 0? 7?55?36.5? 8.6? 0? 5? 2?0? 0? 0? 0? 0? 0? 7? 56?32.6? 9.9? 0? 3? 1?3? 1? 0? 0? 0? 0? 8?57?35.3? 7.4? 0? 3? 3?0? 0? 0? 0? 0? 0? 6? 58?26.2? 12.3? 3? 5? 2?0? 0? 2? 3? 0? 0? 7?59?30.0? 8.6? 0? 4? 1?1? 1? 0? 0? 0? 0? 7? 60?36.1? 11.1? 0? 4? 2?0? 3? 0? 0? 0? 0? 9?61?29.8? 12.3? 0? 3? 0?2? 5? 0? 0? 0? 0? 10? 62?29.7? 13.6? 0? 7? 0?3? 1? 0? 0? 0? 1? 10?63?40.7? 8.6? 0? 3? 0?3? 1? 0? 0? 0? 1? 6 76? Appendix?3.?Characteristics?of?habitat?surrounding?roosts?of?male?Indiana?bats?(Myotis?sodalis),?Pulaski?and?McCreary?counties,?Kentucky,?2001?2003.? Roost?tre? Average?dbh?(cm)?Basal?area?(m?2?/ha)? Pinus? Quercus? Acer? Carya?Other?Number?of?tres?kiled?by?southern?pine?betles? Dead?pines? Living?pines? Dead?non?pines? Living?non?pines? 592?72.5? 4.9? 1? 2? 0? 0? 0? 0? 0? 1? 2? 1?595?19.5? 4.9? 2? 2? 0? 0? 0? 2? 2? 0? 2? 0? 596?29.1? 6.2? 2? 2? 1? 0? 0? 1? 2? 0? 2? 1?800?26.5? 3.7? 1? 1? 0? 0? 1? 0? 2? 0? 1? 0? 887?26.8? 12.3? 0? 7? 1? 0? 2? 0? 0? 0? 0? 0?888?38.4? 7.4? 2? 4? 0? 0? 0? 0? 1? 1? 1? 3? 889?39.8? 9.9? 2? 6? 0? 0? 0? 0? 2? 1? 0? 8?891?20.7? 18.5? 8? 3? 4? 0? 0? 8? 8? 0? 0? 7? 892?39.1? 4.9? 2? 2? 0? 0? 0? 0? 1? 1? 2? 0?894?29.3? 12.3? 8? 2? 0? 0? 0? 2? 5? 2? 0? 2? 895?36.9? 13.6? 7? 2? 0? 0? 2? 6? 7? 0? 1? 3?896?32.2? 11.1? 3? 5? 0? 1? 0? 3? 3? 0? 0? 6? 897?32.4? 14.8? 3? 0? 1? 1? 7? 2? 3? 0? 1? 10?898?37.9? 8.6? 6? 1? 0? 0? 0? 2? 5? 0? 0? 1? 899?40.9? 9.9? 1? 7? 0? 0? 0? 0? 1? 0? 2? 5 77? Appendix?3?continued.? Roost?tre? Average?dbh?(cm)?Basal?area?(m?2?/ha)? Pinus? Quercus? Acer? Carya?Other?Number?of?tres?kiled?by?southern?pine?betles? Dead?pines? Living?pines? Dead?non?pines? Living?non?pines? 992?35.5? 18.5? 9? 2? 2? 0? 2? 8? 9? 0? 0? 6?993?42.4? 9.9? 2? 6? 0? 0? 0? 0? 2? 0? 3? 3? 994?31.7? 6.2? 3? 2? 0? 0? 0? 0? 3? 0? 2? 0?995?36.8? 18.5? 2? 5? 6? 1? 1? 2? 2? 0? 0? 13? 996?28.0? 8.6? 3? 1? 1? 1? 1? 1? 2? 0? 0? 5?997?32.0? 8.6? 1? 3? 1? 1? 1? 2? 2? 0? 0? 5? 999?36.0? 8.6? 7? 0? 0? 0? 0? 4? 7? 0? 0? 0?567?48.1? 1.2? 1? 0? 0? 0? 0? 1? 1? 0? 0? 0? 569?41.4? 6.2? 5? 0? 0? 0? 0? 4? 4? 1? 0? 0?570?20.9? 16.0? 6? 4? 2? 0? 1? 6? 6? 0? 1? 6? 571?43.1? 4.9? 3? 0? 0? 1? 0? 0? 1? 2? 0? 1?576?28.4? 17.3? 7? 2? 2? 2? 1? 7? 7? 0? 0? 7? 674?35.4? 11.1? 3? 3? 3? 0? 0? 0? 1? 2? 0? 8?572?36.5? 7.4? 0? 3? 0? 2? 1? 0? 0? 0? 0? 6? 447?28.8? 17.3? 3? 7? 2? 0? 2? 0? 3? 0? 0? 11?448?30.7? 21.0? 9? 0? 0? 0? 8? 5? 4? 0? 1? 7 78? Appendix?3?continued.? Roost?tre? Average?dbh?(cm)?Basal?area?(m?2?/ha)? Pinus? Quercus? Acer? Carya?Other?Number?of?tres?kiled?by?southern?pine?betles? Dead?pines? Living?pines? Dead?non?pines? Living?non?pines? 446?29.4? 11.1? 4? 0? 0? 0? 5? 1? 3? 1? 0? 6?885?26.3? 3.7? 1? 1? 0? 0? 1? 0? 0? 1? 2? 0? 577?28.2? 8.6? 4? 1? 2? 0? 0? 4? 4? 0? 0? 3?883?29.4? 13.6? 5? 3? 2? 0? 1? 5? 5? 0? 1? 5? 769?38.8? 11.1? 0? 5? 0? 2? 2? 0? 0? 0? 2? 7?583?41.1? 3.7? 3? 0? 0? 0? 0? 3? 3? 0? 0? 0? 581?36.5? 11.1? 5? 4? 0? 0? 0? 4? 5? 0? 1? 3?582?37.4? 12.3? 4? 4? 1? 0? 1? 4? 4? 0? 0? 6? 584?27.7? 1.2? 0? 1? 0? 0? 0? 0? 0? 0? 1? 0?580?29.6? 17.3? 5? 4? 5? 0? 0? 2? 5? 0? 1? 8? 578?29.4? 17.3? 0? 11?1? 0? 2? 0? 0? 0? 2? 12?566?40.7? 14.8? 0? 2? 2? 4? 4? 0? 0? 0? 2? 10? 579?28.4? 9.9? 3? 0? 1? 1? 3? 0? 3? 0? 0? 5?991?31.9? 11.1? 1? 0? 1? 4? 3? 0? 0? 1? 0? 8? 563?35.5? 7.4? 4? 4? 0? 0? 0? 1? 4? 0? 0? 2?562?29.6? 6.2? 4? 1? 0? 0? 0? 0? 4? 0? 1? 0 79? Appendix?3?continued.? Roost?tre? Average?dbh?(cm)?Basal?area?(m?2?/ha)? Pinus? Quercus? Acer? Carya?Other?Number?of?tres?kiled?by?southern?pine?betles? Dead?pines? Living?pines? Dead?non?pines? Living?non?pines? 684?36.0? 3.7? 3? 0? 0? 0? 0? 0? 3? 0? 0? 0?561?32.4? 13.6? 5? 2? 1? 0? 3? 5? 5? 0? 1? 5? 451?24.3? 12.3? 3? 2? 3? 1? 1? 3? 3? 0? 1? 6?990?24.9? 8.6? 3? 2? 2? 0? 0? 3? 3? 0? 0? 4? 989?28.0? 17.3? 7? 0? 5? 0? 2? 7? 7? 0? 0? 7?988?31.0? 7.4? 1? 4? 1? 0? 0? 1? 1? 0? 0? 0? 986?18.2? 14.8? 8? 3? 0? 0? 1? 8? 8? 0? 1? 3?985?41.2? 4.9? 2? 2? 0? 0? 0? 2? 2? 0? 0? 2? 984?26.5? 8.6? 2? 0? 3? 0? 2? 2? 2? 0? 1? 4?983?30.7? 16.0? 7? 2? 3? 0? 1? 6? 7? 0? 0? 6? 458?38.5? 4.9? 3? 1? 0? 0? 0? 0? 1? 2? 1? 0?982?21.5? 11.1? 6? 0? 0? 0? 3? 6? 6? 0? 0? 3? 979?34.7? 13.6? 2? 4? 4? 0? 1? 2? 2? 0? 1? 8?980?32.0? 4.9? 3? 1? 0? 0? 0? 3? 3? 0? 1? 0? 981?33.1? 7.4? 5? 1? 0? 0? 0? 2? 5? 0? 1? 0 80? Appendix?4.?Suitabilityover?time?of?a?subsample?of?trees?used?as?roosts?by?male?Indiana?bats?(Myotis?sodalis),?Pulaski?and?McCreary?counties,?Kentucky,?as?determined?by?revisiting?trees?first?located?during?this?study?and?by?Gumbert?et?al.?(2002).? Datelocated?Roost?tre? October?2001?July?2002?September?2002?April?2003?October?2003?April?2004?Months?a?01?Oct?97?605? Unsuitable? 72?01?Oct?97?606? Unsuitable? 72? 01?Apr?98?123? Unsuitable? 66?01?Apr?98?125? Unsuitable? 66?01?Jul?98?130? Unsuitable? 65? 01?Jul?98?136? Unsuitable? 63?01?Jul?98?146? Unsuitable? 63?01?Oct?98?186? Unsuitable? 60? 01?Oct?98?191? Unsuitable? 62?01?Oct?98?722? Unsuitable? 72?01?Oct?98?732? Suitable? Suitable? Suitable? 72? 01?Oct?98?750? Suitable? 60?01?Oct?98?789? Suitable? 72?01?Apr?99?690? Unsuitable? 54? 01?Jul?99?653? Unsuitable? 45?01?Jul?99?654?Suitable? Unsuitable? 51?01?Oct?99?634? Suitable? Suitable? 50? 01?Oct?99?650? Unsuitable? 50?01?Oct?99?659? Unsuitable? 60?01?Oct?99?662? Unsuitable? 48 81? Appendix?4?continued.?Date?located?Roost?tre? October?2001?July?2002?September?2002?April?2003?October?2003?April2004?Months?a? 29?Sep?01?895?Suitable? Unsuitable? 29?30?Sep?01?891?Suitable? Suitable? 17?02?Oct?01?992?Suitable? Unsuitable? 17? 04?Oct?01?898?Suitable? Unsuitable? 17?04?Oct?01?999?Suitable? Suitable? 29?05?Oct?01?993?Suitable? Unsuitable? 17? 07?Oct?01?996?Suitable? Unsuitable? 17?11?Oct?01?598?Suitable? Suitable? 29?13?Oct?01?888?Suitable? Suitable? Suitable? 29? 14?Oct?01?594?Suitable? Unsuitable? 17?02?Jul?02?575? Suitable? Suitable? 21?03?Jul?02?567? Suitable? Unsuitable? 15? 04?Jul?02?674? Suitable? Suitable? 9?06?Jul?02?570? Suitable? Unsuitable? 16?12?Sep?02?458? Suitable? Suitable? Suitable? 19? 13?Sep?02?459? Suitable? Suitable? 13?14?Sep?02?455? Suitable? Suitable? 13?15?Sep?02?456? Suitable? Suitable? 13? 16?Sep?02?457? Suitable? Suitable? Unsuitable? 19?17?Sep?02?441? Suitable? Suitable? Suitable? Suitable? 19?17?Sep?02?483? Suitable? Suitable? Suitable? 19? 13?Apr?03?446? Suitable? Suitable? 6 82? Appendix?4?continued.?Date?located?Roost?tre? October?2001?July?2002?September?2002?April?2003?October?2003?April?2004?Months?a? 13?Apr?03?883? Suitable? Suitable? Suitable? 12?15?Apr?03?583? Suitable? Suitable? 12?18?Apr?03?582? Suitable? Suitable? 12? 23?Apr?03?566? Suitable? Suitable? 12?04?Oct?03?563? Suitable? Suitable? 6?04?Oct?03?564? Suitable? Suitable? 6? 05?Oct?03?562? Suitable? Suitable? 6?06?Oct?03?451? Suitable? Unsuitable? 6?06?Oct?03?561? Suitable? Suitable? 6? 10?Oct?03?988? Suitable? Suitable? 6?10?Oct?03?989? Suitable? Unsuitable? 6?11?Oct?03?984? Suitable? <1? b?11?Oct?03?986? Suitable? Suitable? 6?12?Oct?03?982? Suitable? Suitable? 6? 12?Oct?03?983? Suitable? Suitable? 6?15?Oct?03?979? Suitable? Suitable? 6?a?Time?from?date?a?bat?was?first?observed?using?a?roost?tree?untilthe?tree?was?observed?unsuitable,?or?until?the?most?recent?date?it?was? observed?suitable?b?Tree?984?became?unsuitable?4?days?after?it?was?used?by?a?bat