Bat$habitat$use$of$restored$jarrah$eucalypt$forests$in$south5western$ Australia.$ ! ! ! $ Joanna$M.$Burgar$ $ $ $ $ School$of$Veterinary$&$Life$Sciences$ Murdoch$University,$Perth,$Western$Australia$ $ $ $ $ Submitted$in$fulfilment$of$the$requirements$for$the$degree$of$ Doctor$of$Philosophy$ $ $ $ $ October$2014$ ! ! ! ! ! Statement'of'Originality! ! ! ! ! I!declare!that!this!thesis!is!my!own!account!of!my!research!and!contains!as!its!main! content!work!that!has!not!previously!been!submitted!for!a!degree!at!any!tertiary! education!institution.! ! ! __________! Joanna!Burgar! ! ! ! i! ! ! ii! ! ! Abstract! Restoration!is!an!important!tool!in!conserving!biodiversity,!yet!passive!faunal! recolonisation!may!take!decades,!or!longer,!to!occur.!This!is!of!particular! conservation!importance!in!biodiversity!hotspots,!such!as!south@western!Australia,! which!are!experiencing!increasing!fragmentation!and!rapidly!drying!climates.! Within!this!hotspot,!I!investigated!the!response!of!nine!insectivorous!tree@dwelling! bat!species!to!restored!mine@pits!in!jarrah!(Eucalyptus*marginata)!forests.!I! assessed!bat!activity!in!restored,!relative!to!unmined,!forests!and!the!suitability!of! restoration!as!foraging!and!roosting!habitat.!Bat!echolocation!call!surveys! measured!bat!activity!in!varying!ages!of!restoration!and!unmined!forest!across!two! years!(2010@2012)!during!both!maternity!and!mating!seasons.!Although!all!bat! species!were!detected!in!both!forest!types,!restored!mine@pits!of!all!ages!had! significantly!different!bat!communities!and!lower!overall!activity!compared!to! unmined!forest.!Habitat!filters!to!bat!use!of!restoration!were!evident!for!the!more! manoeuvrable!bat!species!and!were!predominantly!related!to!midstorey!forest! structure.!Tree!density!was!the!most!important!predictor!of!bat!use!of!restoration! for!less!manoeuvrable!bat!species.!To!determine!the!suitability!of!restored!forest! as!foraging!habitat!I!investigated!the!diet!of!three!species!(Chalinolobus*gouldii,! Nyctophilus*gouldi*and!Vespadelus*regulus)!over!maternity!and!mating!seasons! (2010/2011)!by!examining!prey!remains!in!faecal!samples.!I!used!high@throughput! sequencing!and!bioinformatics!analyses!to!phylogenetically!group!prey!DNA!and! found!that!niche!partitioning!occurred,!with!dietary!divergence!positively!related! ! iii! to!bat!ecomorphological!divergence.!In!addition,!I!assessed!the!foraging!potential! of!restored!forest!and!found!that!prey!occurrence!did!not!necessarily!equate!to! prey!accessibility!for!all!bat!species.!There!was!a!synergistic!effect!of!vegetation! structure!and!insect!biomass!for!edge!foraging!bat!species.!To!determine!the! suitability!of!restoration!as!roosting!habitat!I!used!telemetry!to!radio@track!36!bats! from!two!species!(N.*gouldi*and!V.*regulus)!to!59!distinct!roosts.!Not!one!bat!was! found!roosting!in!restored!forest!and!individuals!preferred!roosting!in!mature,!tall! trees!in!intermediate!to!late!stages!of!decay.!My!research!clearly!shows!that! restored!forest!does!not!yet!provide!suitable!foraging!or!roosting!habitat!for!all! jarrah!forest!bats.!Improving!habitat!suitability!through!management! manipulations,!such!as!thinning!and!burning,!may!accelerate!bat!recolonisation!of! restored!forest.!In!the!interim,!retention!of!mature!forest!patches!is!necessary!for! conserving!and!maintaining!bat!populations!across!restored!landscapes.! ! ! iv! ! Table&of&Contents! STATEMENT$OF$ORIGINALITY$ I! ABSTRACT$ III! TABLE$OF$CONTENTS$ V! ACKNOWLEDGEMENTS$ 2! CHAPTER$ONE:$GENERAL$INTRODUCTION$ 5! STUDY$AREA$&$RESTORATION$PROCESS$ 7! BATS$OF$THE$NORTHERN$JARRAH$FOREST$ 9! EFFECTS$OF$DISTURBANCE$ON$BATS$ 17! HYPOTHESES$FOR$BAT$USE$OF$RESTORATION$ 25! THESIS$CONTENT$ 26! CHAPTER$TWO$ 29! INTRODUCTION$ 29! METHODS$&$MATERIALS$ 34! RESULTS$ 43! DISCUSSION$ 55! CHAPTER$THREE$ 66! ABSTRACT$ 66! INTRODUCTION$ 67! METHODS$&$MATERIALS$ 72! RESULTS$ 78! DISCUSSION$ 89! CHAPTER$FOUR$ 97! INTRODUCTION$ 97! METHODS$&$MATERIALS$ 102! RESULTS$ 115! DISCUSSION$ 126! CHAPTER$FIVE$ 136! INTRODUCTION$ 136! METHODS$&$MATERIALS$ 140! RESULTS$ 151! DISCUSSION$ 165! CHAPTER$SIX:$GENERAL$DISCUSSION$ 175! APPENDIX$ONE:$BAT$CALL$AUTOMATION$ 193! INTRODUCTION$ 193! MATERIALS$AND$METHODS$ 196! RESULTS$ 210! DISCUSSION$ 219! ! v! APPENDIX(TWO:(SUPPLEMENTARY(FIGURES(&(TABLES( 224! CHAPTER(FOUR( 224! CHAPTER(FIVE( 226! APPENDIX(ONE( 234! LITERATURE(CITED( 238! vi! ! ! ! Acknowledgements! This!thesis!arose!out!of!work!undertaken!since!I!came!to!Australia!in!2010.!This! work!would!not!have!been!possible!without!the!help!and!support!of!a!great!many! people!along!the!way.!I!would!never!have!been!inspired!to!undertake!a!PhD! without!the!two!best!role!models!a!daughter!could!hope!for,!my!parents,!who!both! taught!and!never!asked!if,!but!instead!where,!I!was!going!to!university.!To!my! father,!who!helped!me!with!my!science!homework!and!gave!me!my!aptitude!for! math,!I!am!so!fortunate!to!have!had!you!for!a!dad!and!I!only!wish!you!could!be!here! to!see!me!graduate!–!you!are!in!my!heart!always.!To!my!mother,!who!listened!to!all! of!my!challenges!and!struggles!with!fieldwork,!analyses!and!writing,!and!helped! with!grammar!to!the!bitter!end,!you!are!an!inspiration!for!tackling!a!PhD!while! working!and!yet!still!managing!to!ensure!a!loving!home!for!our!family!–!I!aspire!to! be!as!selfless!and!generous!as!you.!To!my!sister,!who!gave!me!unwavering!support,! always!managed!to!find!me!work!and!thankfully!even!a!job!that!gave!me!a!good! start!in!stats,!thank!you!for!being!so!close!even!when!you!were!half!a!world!away.! ! I!am!forever!indebted!to!my!two!supervisors,!Dr.!Michael!Craig!and!Dr.!Vicki! Stokes,!without!whose!unending!assistance!this!thesis!would!not!have!been! possible.!I!am!also!extremely!grateful!to!Dr.!Kyle!Armstrong!for!lending!me!field! equipment,!advising!me!on!bat!call!identification,!and!providing!reference!bat!calls;! to!Dr.!Allan!Wills!for!the!loan!of!light!traps!and!assistance!with!invertebrate! identification;!to!Dr.!Paul!Webala!for!giving!me!a!quick!introduction!to!jarrah! forest!bats,!answering!countless!emails!thereafter,!and!providing!reference!bat! calls;!and!to!Sian!Contarino!for!persevering!and!assisting!with!sorting!thousands!of! invertebrate!samples.!I!am!also!grateful!to!the!many!Murdoch!and!University!of! Western!Australia!undergraduate!students!who!came!out!in!the!field!or!spent!a! day!in!the!lab!to!assist!me.!! ! I!will!also!be!forever!grateful!to!Richard!Hobbs!and!the!entire!Ecosystem! Restoration!&!Intervention!Ecology!Research!Group!(ERIE)!at!the!University!of! Western!Australia!for!being!such!an!inclusive!lab!group,!both!intellectually!and! 2! ! socially.!Having!the!opportunity!to!present!ideas!and!rehearse!conference!papers! in!a!non@threatening!and!collegial!environment!certainly!improved!my!public! speaking!confidence,!and!perhaps!was!the!reason!I!was!able!to!take!runner@up! student!presenter!at!my!last!conference!as!a!student.!Friday!afternoon!(sometimes! progressing!into!night)!beers!were!also!a!great!way!to!let!off!some!of!the!work@ week!stress,!and!make!weekend!working!that!much!more!enjoyable.! ! I!would!also!like!to!thank!all!of!the!people!who!made!Australia!feel!like!home!for! me!and!kept!me!inspired!during!what,!at!times,!felt!like!a!never@ending!journey.!I! don’t!know!if!I!would!have!made!it!through!the!day!without!the!morning!coffee! breaks!with!the!MUCRU!group,!who!made!special!concessions!to!include!me!as! another!echolocating!animal!researcher.!To!the!inspiring!Krista!Nicholson!for! reminding!me!of!the!importance!of!quality!data!collection!as!well!as!being!a! wonderful!distraction!and!the!perfect!person!to!have!sitting!just!behind!me.!To! Jenny!Smith!for!keeping!me!sane!with!yoga,!tea!study!dates,!and!sustainability,!and! for!reminding!me!of!good!ole!North!America!with!your!unbelievable!recollection!of! pop@culture!while!making!such!witty,!intelligent!remarks.!To!Emily!Fisher!for! joining!me!in!exploration!of!south@west!surf!spots,!keeping!me!company!at! Murdoch!on!those!weekend!work!days,!and!for!being!such!a!sweet!and! understanding!friend.!To!Simon!Allen!for!motivating!me!to!get!my!PhD!submitted! on!time…come!to!think!of!it,!am!I!not!owed!a!dinner,!or!did!we!resolve!to!keep!it!as! a!gentleman’s!bet?! ! To!Maggie!Triska!for!sharing!in!the!fun!of!dozens!of!veg!surveys,!chit@chatting!PhD! issues!during!long!drives!to!and!from!Huntly,!being!a!stats!guru!and!discussing! stats!even!when!we!were!supposed!to!be!out!for!fun,!being!my!wildlife!spotting! partner!in!crime,!but!most!importantly!for!being!that!wonderful!ray!of!North! Dakota!sunshine!that!I!was!lucky!enough!to!pick!up!from!the!airport!on!your!first! day!in!town!and!being!such!a!good!friend,!one!that!I!hope!to!never!lose.!To! Christine!Allen!for!reminding!me!of!the!beauty!of!the!south@west!with!camping! trips!and!writing!retreats,!being!another!odd!person!in!the!drive!for!sustainability! and!wacky!food!choices!and,!of!course,!for!all!the!fun!dancing!!To!Bridget!Johnson! for!the!absolute!best!Ridgefield!experience,!sharing!in!outrageous!silliness,!making! ! 3! me!laugh,!and!generously!giving!me!a!place!to!stay!and!a!car!to!use!when!I!was! without.!!And!of!course!to!the!(in)famous!Dr.!Jodi!Price!for!joining!me!at!punk! shows!(not!enough,!though!),!making!me!loads!of!yummy!vegan!meals!and!treats,! putting!together!a!nearly@there!PhD!package!for!the!last!push,!sharing!your!office! with!me,!and!always!being!up!for!having!a!good!time.! ! I!acknowledge!the!support!of!Murdoch!University!for!funding!my!scholarship!and,! through!the!Murdoch!University!Animal!Ethics!Committee,!providing!the!approval!