MIAMI$UNIVERSITY$ The$Graduate$School$ $ $ Certificate)for)Approving)the)Dissertation) ) We)hereby)approve)the)Dissertation) $ of) ) Jessica)René)Peebles;Spencer) ) Candidate$for$the$Degree$ ) ) Doctor)of)Philosophy) $ $ ______$ David)L.)Gorchov,)Director) $ ______$ M.)Henry)H.)Stevens,)Reader) $ ______$ Melany)C.)Fisk,)Reader) ) ______) R.)James)Hickey,)Reader) ) ______) Thomas)O.)Crist,)Graduate)School)Representative) ) ) ABSTRACT) ) ) ) INTERACTIONS$BETWEEN$INVASIVE$SHRUB,$LONICERA)MAACKII,$$ AND$A$GENERALIST$HERBIVORE,)WHITEETAILED$DEER,$ $IN$SOUTHWESTERN$OHIO$FORESTS) ) ) by)Jessica)R.)Peebles;Spencer) ) ) While)invasive)plants)and)generalist)herbivores)have)negative)direct)effects)on)plant) communities,)their)indirect)and)interactive)effects)have)received)far)less)focus.) I)address)the)effects)of)an)abundant)herbivore,)White;tailed)deer)(Odocoileus* virginanus),)and)the)invasive)shrub)Amur)honeysuckle)(Lonicera*maackii))on)forest)plant) communities,)focusing)on)determining)whether)their)interactions)are)synergistic,) antagonistic,)or)additive.))Within)the)Miami)University)Natural)Areas,)I)sampled)plant) species)richness)and)cover)in)five)20)x)20)m2)deer)exclosures)and)paired)controls,)each)with) L.*maackii*removed)from)half)of)the)plot.)I)found)antagonistic)interactions)of)L.*maackii*and) deer)on)the)tree)seedling)cover,)suggesting)that)L.*maackii*mitigates)the)negative)effect)of) deer)on)trees,)facilitating)tree)survival)and)growth.))) To)investigate)whether)L.*maackii*provides)a)refuge)for)tree)seedlings)from)deer) browse,)I)planted)tree)seedlings)under)and)away)from)the)L.*maackii*canopy,*with)and) without)deer)exclosures,)and)measured)their)growth,)survival)and)browse)status.))I)found) facilitation)by)L.*maackii*on)survival)and)leaf)count)of)Acer*saccharum)seedlings:)where) deer)had)access,)seedlings)planted)under)L.*maackii*had)higher)survival)and)more)leaves.)) This)suggests)that)L.*maackii)serves)as)a)refuge)for)tree)seedlings)from)deer)browse.) Finally,)I)explored)the)direct)effects)of)deer)on)L.*maackii,*addressing)whether)deer) browse)on)L.*maackii*is)sufficient)to)reduce)its)cover.))In)the)20)x)20)m2)plots)I)scored)leaf) frequency)of)L.*maackii*in)the)height)ranges)browsed)by)deer,)as)well)as)basal)area)(BA))of) all)stems)of)L.*maackii.))I)found)access)to)deer)strongly)reduced)L.*maackii)leaf)frequency) within)the)height)range)of)deer)browse,)and)modestly)reduced)BA)growth.)This)suggests) that)deer)affect)the)architecture)of)L.*maackii)more)than)its)growth) These)results)indicate)that)considering)interactive)effects)is)fundamental)to) understanding)the)roles)of)these)factors)in)complex)natural)systems.)I)suggest)that) management)for)tree)regeneration)should)focus)first)on)reducing)deer)density,)and)later)on) removal)of)L.*maackii.)) ) INTERACTIONS)BETWEEN)INVASIVE)SHRUB,)LONICERA*MAACKII,)) AND)A)GENERALIST)HERBIVORE,)WHITE;TAILED)DEER,) )IN)SOUTHWESTERN)OHIO)FORESTS) ) )
A$DISSERTATION$
)
Submitted)to)the)Faculty)of))
Miami)University)in)partial))
fulfillment)of)the)requirements))
for)the)degree)of))
Doctor)of)Philosophy))
Department)of)Biology)
Ecology,)Evolution)and)Environmental)Biology)
)
by)
)
Jessica)René)Peebles;Spencer)
)
Miami)University)
Oxford,)Ohio))
)
2016)
)
Dissertation)Director:)David)L.)Gorchov) Table$of$Contents$ FRONT$MATTER$ LIST)OF)TABLES)...... )IV) LIST)OF)FIGURES)...... )V) DEDICATION)...... )VI) ACKNOWLEDGEMENTS)...... )VII) CHAPTER$1.$INTERACTIONS$BETWEEN$INVASIVE$SHRUB,$LONICERA)MAACKII,$AND$A$ GENERALIST$HERBIVORE,$WHITEETAILED$DEER,$IN$SOUTHWESTERN$OHIO$FORESTS$...... $1) REFERENCES)...... )6) CHAPTER$2.$EFFECTS$OF$AN$INVASIVE$SHRUB,$LONICERA)MAACKII,$AND$A$GENERALIST$ HERBIVORE,$WHITEETAILED$DEER,$ON$FOREST$FLOOR$PLANT$COMMUNITY$COMPOSITION$11) INTRODUCTION)...... )11) METHODS)...... )15) Field*Methods*...... *15) Data*analysis*...... *16) RESULTS)...... )18) Species*Richness*...... *18) Species*Composition*...... *19) Indicator*Species*...... *19) Cover*of*Functional*Types*...... *19) Bare*Ground*...... *20) Invasive*species*...... *20) Herb*reproduction*...... *20) DISCUSSION)...... )21) Lonicera*maackii*direct*effects*...... *21) Deer*Direct*Effects*...... *22) Interactions*...... *23) Support*for*Hypotheses*...... *24) REFERENCES)...... )26) CHAPTER$3.$ARE$NATIVE$TREE$SEEDLINGS$FACILITATED$BY$AN$INVASIVE$SHRUB$WHERE$ WHITEETAILED$DEER$ARE$ABUNDANT?$...... $46) INTRODUCTION)...... )46) MATERIALS)AND)METHODS)...... )48) Field*Methods*...... *48) Data*Analysis*...... *49) RESULTS)...... )51) Survival*...... *51) Browse*status*...... *51) Final*Leaf*Count*...... *51) Change*in*Height*...... *52) Root*Mass,*Shoot*Mass*and*Root:Shoot*Ratio*...... *52) DISCUSSION)...... )52) Facilitation*...... *52) Competition*...... *53) Additional*Findings*...... *53) REFERENCES)...... )56)
) ii) CHAPTER$4.$BROWSE$BY$WHITEETAILED$DEER$DECREASES$COVER$AND$GROWTH$OF$THE$ INVASIVE$SHRUB,$LONICERA)MAACKII$...... $73) INTRODUCTION)...... )73) METHODS)...... )75) Field*Methods*...... *75) Data*Analysis*...... *76) RESULTS)...... )76) DISCUSSION)...... )76) REFERENCES)...... )79) CHAPTER$5.$CONCLUSIONS$...... $87) GOALS)...... )87) NON;ADDITIVE,)ANTAGONISTIC)INTERACTIVE)EFFECTS)ON)PLANT)COMMUNITY)COMPOSITION)...... )87) FACILITATIVE)INTERACTIONS)BETWEEN)L.*MAACKII*AND)TREE)SEEDLINGS)...... )88) DECREASED)GROWTH)AND)COVER)OF)L.*MAACKII*IN)THE)PRESENCE)OF)DEER)...... )88) SYNTHESIS)...... )89) APPENDIX$...... $91) ADDITIONAL)TABLES)AND)FIGURES)FROM)CHAPTER)2)...... )91) ADDITIONAL)TABLES)AND)FIGURES)FROM)CHAPTER)3)...... )111) ADDITIONAL)TABLE)FROM)CHAPTER)4)...... )119) ) ) )
) iii) List$of$Tables$ Chapter)2.)) ) Table)1.)Repeated)measures)ANOVA)table)for)plant)species)richness)...... )33) Table)2.)Significance)of)treatment)effects)on)species)composition)of)forest)floor)plants)by) year)and)season)...... )34) Table)3.)Indicator)species)of)significant)compositional)treatment)effects)...... )35) Table)4.)Significance)of)treatment)effects)on)area)of)bare)ground)by)year)and)season.)...... )36)) ) Chapter)3.)) $ Table)1.)ACSA)seedling)survival)by)treatment)type.)...... )61) Table)2.)Significance)of)treatment)effects)on)leaf)count)of)surviving)seedlings)of)each) species.)...... )62) Table)3.)Significance)of)treatment)effects)on)change)in)height)of)surviving)seedlings)of)each) species.)...... )63) ) Chapter)4.)* * Table)1.))Browse)on)first;year)Lonicera*maackii*twigs)in)five)deer)access)plots)within)Miami) University)Natural)Areas.)...... )83) ) ) )
) iv) List$of$Figures$ ) Chapter)2.)) ) Figure)1.))Predictions)of)plant)community)responses)...... )37) Figure)2.))Map)of)study)sites)in)the)Miami)University)Natural)Areas)...... )38) Figure)3.))Schematic)of)study)design)...... )39) Figure)4.))Boxplots)of)spring)and)summer)plant)species)richness)across)the)4)years)of)the) study)...... )40) Figure)5.))Boxplots)of)cover)of)plant)functional)types)by)treatment)in)spring)2012,)2013,) and)2014,)and)summer)2013)...... )41) Figure)6.))Percent)tree)seedling)cover)in)Spring)2013)...... )42) Figure)7.))Percent)tree)seedling)cover)in)Summer)2013)...... )43) Figure)8.))Boxplots)of)percent)bare)ground)in)spring)2013)and)2014)and)summer)2012,) 2013,)and)2014)...... )44) Figure)9.))Mean)number)(±SE))of)reproductive)individuals)of)Sanicula*odorata)and)Alliaria* petiolata*per)plot)by)treatment)type)...... )45) ) Chapter)3.)) ) Figure)1.))Predictions)of)tree)seedling)responses)(survival,)growth,)etc.))...... )64) Figure)2.))Schematic)of)study)design)...... )65) Figure)3.))Map)of)study)locations)within)the)Miami)University)Natural)Areas)...... )66) Figure)4.))Percent)survival)(+/;)SE))of)seedlings)...... )67) Figure)5.)Survival)curves)for)Acer*saccharum)(ACSA))...... )68) Figure)6.))Interaction)plot)of)Acer*saccharum)survival,)mean)±)SE)for)each)treatment)...... )69) Figure)7.))Final)leaf)count)of)seedlings)...... )70) Figure)8.))Interaction)plot)of)Acer*saccharum)seedling)leaf)count,)mean)±)SE)for)each) treatment)...... )71) Figure)9.)Change)in)height)between)July)2014)and)June)2015)of)seedlings)...... )72) ) Chapter)4.* * Figure)1.))Boxplot)of)Lonicera*maackii*basal)area)change)between)2010)and)2015)...... )84) Figure)2.))Mean)frequency)of)Lonicera*maackii*foliage)at)each)of)four)heights)in)deer)access) and)deer)exclosure)plots)...... )85) Figure)3.))Mean)frequency)of)Lonicera*maackii*foliage)at)each)height)range)in)deer)access) and)deer)exclosure)plots)...... )86) ) )
) v) Dedication$ ) For)Grandpa)Dave) ) For)instilling)in)me)a)love)of))"outside")and)) as)an)apology)for)ever)asking)when)the)mall)opened)while)fishing.) $ )
) vi) Acknowledgements$
My)advisor,)Dr.)Dave)Gorchov.)Thank)you)for)believing)I)could)do)it,)for)challenging) me)to)do)it)better,)for)encouraging)me)to)speak)up)before)I)have)finished)mentally)debating) if)I)am)right,)for)valuing)my)opinions,)for)reading)this)way)before)any)human)should)have) been)subjected)to)it,)but)most)of)all,)for)always)being)available)when)I)needed)you.)I)am) honored)to)have)spent)all)this)time)working)with)you.)
My)committee,)Dr.)Tom)Crist,)Dr.)Hank)Stevens,)Dr.)Melany)Fisk)and)Dr.)Jim)Hickey.) Thank)you)for)your)encouragement)and)support)throughout)this)process,)for)your) innumerable)comments,)and)for)always)working)with)my)"abbreviated")timelines.))Your) input)has)had)a)significant)effect)(P)<)0.05).)There)are)significantly)fewer)uses)of)the)word) significant)because)of)the)significant)impact)you)have)had)on)my)writing.)
The)Gorchov)lab,)past)and)present,)the)adoptees)we)picked)up)along)the)way,)and) my)other)grad)school)friends.)Thank)you)for)your)ideas,)for)your)camaraderie,)and)for)your) literal)blood,)sweat,)and)tears.))The)burden)is)lighter)when)it)is)shared.))Jen,)thank)you)for) doing)it)all)first)and)then)guiding)me)through)it.)MB,)thank)you)for)being)my)very)first)and) very)best)friend)at)Miami.)We)had)no)idea)what)we)were)getting)ourselves)into.)
My)community.))Thank)you)for)investing)in)me)over)the)years)and)for)praying)me)to)) and)through)this)point.))I)am)blessed)to)have)you)in)my)life.)
My)immediate)family,)Mom,)Dad,)Gregor,)and)my)Grandparents.))Thank)you)and)I) love)you)are)just)not)enough.))Your)sacrifices)were)unfathomable)and)a)few)lines)on)this) dissertation)do)not)begin)to)address)your)importance)in)my)life.))I)am)who)I)am)because)of) you.)
My)husband.))Bob,)you)believed)in)me)when)I)didn't)believe)in)me.)You)are)the)one) person)in)the)world)who)knows)how)much)of)me)went)into)this.)Thank)you)for)being)strong) for)me.)Your)love)gives)me)courage.))I)thank)God)for)you)every)day.))Forever)and)for)always,) for)real)and)for)true.)Now)what's)our)next)adventure?
) vii) Chapter$1.$Interactions$between$Invasive$Shrub,$Lonicera)maackii,$and$a$ Generalist$Herbivore,$WhiteEtailed$Deer,$in$Southwestern$Ohio$Forests$ )
With)the)ever;increasing)prevalence)of)global)travel)and)trade,)humans)have) purposefully)or)accidentally)introduced)many)species)of)plants)and)animals)to)areas)that) are)outside)of)their)native)ranges)(Mack)et)al.)2000,)Keane)and)Crawley)2002).)))Once)these) organisms)are)introduced,)some)become)disruptive)and)detrimental)to)the)environments) they)invade)(Mack)et)al.))2000).)))Invasive)plant)species)are)introduced)through)many) channels,)such)as)horticulture,)plantings)for)erosion)control,)or)contamination)of)crop) seeds,)among)other)reasons)(Reichard)and)White)2001).))) The)effects)of)invasive)plant)species)on)other)plant)species)have)been)widely) documented,)and)generally)determined)to)be)negative)in)nature)(Mack)et)al.))2000).))) Invasive)woody)species)can)be)particularly)difficult)to)manage,)and)they)can)prevent)the) growth)and)progression)into)the)canopy)of)native)trees)and)may)have)cascading)effects)on) other)biota)(Webster)et)al.)2006).))For)example,)herb)species)richness)was)41%)less)and) woody)stems)were)75%)less)abundant)in)otherwise)similar)areas)with)vs.)without) Ligustrum*sinense)Lour.)shrubs)(Merriam)and)Feil)2002).))High)density)of)the)invasive) shrub)Rhamnus*frangula*Mill.)allowed)for)less)than)10%)of)native)tree)seedlings)to)survive) to)the)point)where)they)can)grow)through)the)canopy)of)the)invasive)(Fagan)and)Peart) 2004),)and)caused)reductions)in)tree)seedling)density,)possibly)by)inhibiting)the) establishment)of)these)seedlings)(Frappier)et)al.)2004).))Tree)seedling)density)and)annual) plant)cover)were)significantly)reduced)within)areas)with)density)of)the)invasive)shrub,) Lonicera*tatarica*L.)(Woods)1993).))) Lonicera*maackii*(Rupr.))Herder,)Amur)honeysuckle,)is)one)of)several)bush) honeysuckle)species)that)are)invasive)in)the)eastern)United)States)(Miller)2006).)))Its)native) range)includes)China,)Japan,)Korea,)and)the)Russian)Far)East,)and)is)named)after)the)Amur) River,)which)forms)the)division)between)Russia)and)China)(Luken)and)Thieret)1996).) Lonicera*maackii)is)a)large)shrub)with)dense,)medium)to)dark)green)foliage)that)grows)to) heights)of)up)to)6)meters)(Luken)and)Thieret)1996);)it)has)been)cultivated)in)gardens)for)its)
) 1) flowers)and)bright)red)fruits)from)as)early)as)the)1880s)in)Europe)(Luken)and)Thieret) 1996).))))It)was)introduced)into)the)United)States)and)established)in)the)Harvard)University) Botanical)Gardens)in)1903)(Luken)and)Thieret)1996).))The)current)distribution)of)L.* maackii)spans)the)eastern)half)of)the)United)States;)L.*maackii)is)considered)invasive)and)is) regulated)in)8)of)those)states)(EDDMapS)2015).))))It)was)widely)available)for)horticultural) planting,)and)subsequently)also)escaped)from)cultivation)(Luken)and)Thieret)1996).)) Lonicera*maackii)seeds)are)primarily)dispersed)by)birds)(Bartuszevige)et)al.)2006),)but)can) also)be)dispersed)by)white;tailed)deer)(Castellano)and)Gorchov)2013,)Guiden)et)al.))2015).))) Lonicera*maackii)shrubs)become)reproductively)active)3;8)years)after)establishment)and)at) >)1)m)in)height,)with)size)being)highly)correlated)with)age)(Deering)and)Vankat)1999,) Gorchov)et)al.)2014).))) ) The)prodigious)success)of)L.*maackii)in)eastern)forests,)attaining)densities)of)up)to) 6800)shrubs)per)hectare)(Trisel)and)Gorchov)1994),)can)be)attributed)to)many)factors.))) Lonicera*maackii)has)an)extended)leaf)phenology,)leafing)out)earlier)than)many)native) plants)and)retaining)those)leaves)long)after)other)deciduous)plants)have)senesced,)allowing) it)extended)time)to)photosynthesize)compared)with)its)competitors)(McEwan)et)al.))2009).))) In)addition,)L.*maackii)has)a)higher)freeze)tolerance)compared)to)other)shrubs,)allowing)it) to)survive)late)spring)freezes)which)more)negatively)impact)its)competitors)(McEwan)et)al.)) 2009).)))Lonicera*maackii)can)opportunistically)establish)in)canopy)gaps)following)forest) disturbance,)and)once)the)gaps)close,)it)can)persist)in)the)closed)canopy)forest)due)to)its) high)shade)tolerance)(Luken)and)Thieret)1996).)))Within)its)introduced)range,)L.*maackii)is) not)affected)by)any)of)its)specialist)herbivores)(Lieurance)and)Cipollini)2012))and)may) therefore)benefit)from)enemy)release)(Keane)and)Crawley)2002).))I)provide)a)more) thorough)discussion)of)L.*maackii*and)the)Enemy)Release)Hypothesis)in)chapter)4.*These) characteristics)have)allowed)L.*maackii)to)become)dominant)in)the)communities)it)invades,) decreases)growth,)survival,)density)and)cover)of)tree)seedlings)and)saplings)(Collier)et)al.) 2002,)Gorchov)and)Trisel)2003,)Hartman)and)McCarthy)2008).))The)effects)of)L.*maackii* invasion)on)forest)floor)plant)communities)are)explored)as)part)of)chapter)2.) ) While)the)direct)effects)of)invasive)shrubs)on)forest)floor)plants,)such)as)tree) seedlings,)are)well)studied,)and)generally)considered)to)be)negative)(Woods)1993,)Merriam) and)Feil)2002,)Fagan)and)Peart)2004,)Frappier)et)al.)2004,)Webster)et)al.)2006),)there)are)
) 2) some)situations)in)which)these)effects)may)not)so)straightforward.))In)stressful) environments,)such)as)those)caused)by)severe)water)limitation)(Bertness)and)Callaway) 1994))or)intense)herbivore)browse)pressure)(Bertness)and)Callaway)1994,)Gómez;Aparicio) et)al.)2008),)plants)may)instead)interact)in)a)positive)manner,)though)amelioration)of) physical)abiotic)conditions)(Callaway)1995).))This)interaction)is)an)example)of)‘facilitation,’) a)positive)interaction)that)is)not)consumptive)in)nature)and)that)benefits)at)least)one)of)the) species)involved)(Merriam)and)Feil)2002).))When)the)stressor)is)water)limitation,)this) facilitation)is)exemplified)by)one)plant,)usually)called)a)“nurse)shrub,”)providing)a)shady) environment)which)reduces)evaporation,)allowing)for)another)plant)species)to)benefit)from) the)less)arid)microhabitat)(Bertness)and)Callaway)1994),)When)the)stressor)is)intense) herbivore)browse,)this)facilitation)is)exemplified)by)a)well)defended)or)low)browse) preference)shrub)species)that)is)avoided)by)herbivores,)allowing)other)plant)species)to) benefit)from)the)lessened)browse)pressure)under)its)canopy)(García)and)Obeso)2003,)Graff) et)al.)2007,)Smit)et)al.)2007,)Vandenberghe)et)al.)2009).))In)my)third)chapter,)I)argue)that) the)current)herbivore)pressure)in)my)study)area)is)sufficient)to)be)considered)a)stressful) environment)for)plants,)and)I)will)address)whether)by)nature)of)its)abundance)and) relatively)low)herbivore)preference,)if)L.*maackii*can)function)as)a)refuge)from)browse)for) tree)seedlings,)facilitating)their)growth)and)survival.))) The)range)of)L.*maackii*throughout)North)America)overlaps)with)that)of)White; tailed)deer,)Odocoileus*virginanus*Zimmerman,)an)overabundant)generalist)herbivore)that) exhibits)this)intense)browse)pressure)(Hirth)1977,)Russell)et)al.)2001,)Gallina)and)Lopez) Arevalo)2008,)EDDMapS)2015).))Within)the)United)States,)White;tailed)deer)faced)a) massive)population)decline)in)the)middle)of)the)nineteenth)century)(Côté)et)al.)2004).)))In) response,)many)protective)measures,)such)as)game)laws)and)restricted)hunting)seasons,) were)put)into)place)(Côté)et)al.)2004).)))Additionally,)the)disappearance)of)the)wolf)and)the) mountain)lion,)and)increasing)availability)of)forage)in)the)form)of)high)density) monocultures)of)corn)and)soybeans,)lead)to)the)density)rebounding)(Sotala)and)Kirkpatrick) 1973,)Côté)et)al.))2004).)))Estimates)of)pre;settlement)deer)densities)range)from)2;4) deer/km2)across)their)entire)North)American)range)(Rooney)2001),)and)have)increased)to) average)densities)greater)than)10)deer)per)square)kilometer)(Rooney)and)Waller)2003).)))) White)tailed)deer)often)reach)densities)30;50)deer/km2)where)not)managed)(McShea)
) 3) 2012),)though)in)one)site)(Sharon)Woods)Metro)Park,)Ohio))densities)as)high)as)over)100) deer/km2)were)reached)and)noted)as)the)cause)of)local)extirpations)of)more)than)150) vascular)plant)species)in)that)site)(Peck)and)Stahl)1997).) The)overabundance)of)White;tailed)deer)can)cause)negative)impacts)through)both) direct)and)indirect)effects)(Rooney)and)Waller)2003))even)at)low)to)moderate)densities,)3; 10)deer/km2)(Horsley)et)al.)2003,)Côté)et)al.)2004,)Rooney)2009).))Their)negative)effects)are) both)direct,)including)decreased)crop)yields)(Palmer)et)al.))1982))and)browse)on) economically)and)ecologically)important)tree)species)(Horsley)et)al.))2003,)Côté)et)al.)) 2004),))as)well)as)indirect,)including)increases)in)tick;borne)illnesses)in)humans)(Bloemer) et)al.)1986,)Allan)et)al.)2010).))The)effects)of)overabundant)deer)on)forest)floor)plant) communities)are)explored)in)chapter)2.) While)invasive)plants)and)generalist)herbivores)have)each)been)shown)to)have) direct)negative)effects)on)plant)communities,)their)indirect)and)interactive)effects,)often) fundamental)to)understanding)invasions)and)their)impacts)on)complex)ecosystems,)have) received)far)less)focus,)at)least)in)part)due)to)the)difficulty)of)isolating)and)assessing)these) interactions)(Strauss)1991,)Wootton)1994a,)1994b,)2002,)Krivtsov)2004,)White)et)al.)2006).) There)have)been)few)studies)simultaneously)addressing)the)impacts)of)both)L.*maackii*and) deer)on)the)forest)floor)plant)community.))Of)those)that)have,)two)have)suggested) interactive)effects)as)a)result)of)their)findings)(Gorchov)and)Trisel)2003,)Cipollini)et)al.) 2009))and)two)have)explicitly)tested)for)interactive)effects,)one)finding)them)on)annuals,) spring)perennials)and)on)leaf)count)of)a)herb)that)was)very)favored)by)deer)(Christopher)et) al.)2014),)and)one)finding)no)interactions)between)any)mammalian)consumer)(including) deer))on)native)or)invasive)plant)species)abundance)or)richness)(Orrock)et)al.)2015).)*)In)my) second)chapter)I)explicitly)address)the)indirect)effects)of)deer)and)L.*maackii*on)the)forest) floor)plant)community,)particularly)in)terms)of)plant)richness)and)community)composition,) focusing)on)determining)whether)these)effects)are)synergistic);)with)combined)effects)being) more)negative)for)the)plant)community)than)either)individually,)antagonistic);)with) combined)effects)being)less)negative,)or)additive);)with)combined)effects)being)the)just)the) sum)of)the)negative)effects)of)each)factor.) The)direct)impacts)of)deer)on)L.*maackii*are)also)relatively)unknown.)While)the) browse)preference)by)deer)of)L.*maackii*is)not)well)studied,)its)leaves)have)low)palatability)
) 4) for)insects)(Lieurance)and)Cipollini)2012,)2013),))and)the)congeneric)invasive)shrub)L.* morrowii*A.)Gray,*is)of)moderate)preference)to)deer)(Averill)et)al.)2016)).))Additionally,) congeneric)invasive)vine,)L.*japonica*Thunb.,)has)long)been)planted)as)deer)forage) (Stransky)1984).)))Deer)may)be)important)in)the)long)distance)dispersal)of)seeds)through) ingestion)(endozoochory))of)L.*maackii)(Castellano)and)Gorchov)2013,)Guiden)et)al.))2015).) Deer)browse)on)L.*maackii*in)areas)of)several)different)vegetation)types)in)the)Miami) University)Natural)Areas)in)southwestern)Ohio,)with)the)percentage)of)first;year)twigs) browsed)between)leaf)flush)and)late)May)ranging)from)3.2%)in)red)cedar)forests)to)5.5%)in) forest)edge)habitats,)with)4.9%)of)twigs)browsed)in)deciduous)forest)(K.)Martinod,) unpublished)data).)))In)my)fourth)chapter,)I)will)address)the)direct)effects)of)deer)on)L.* maackii,*investigating)whether)the)amount)of)browse)by)deer)on)the)canopy)of)L.*maackii*is) sufficient)to)reduce)its)cover.)) This)dissertation)addresses)how)these)two)factors,)a)shrub)layer)of)the)invasive)L.* maackii*and)overabundant)deer,)affect)forest)communities)in)southwestern)Ohio.)I’ll) address)both)the)direct)effects)of)deer)on)L.*maackii*(chapter)4),)and)the)indirect)effects)of) deer)and)L.*maackii*on)forest)floor)species)composition)and)richness)(chapter)2))as)well)as) specific)impacts)on)tree)seedlings)(chapter)3).)Taken)together,)these)studies)will)further)the) knowledge)of)the)interactions)between)native)and)invasive)species,)and)how)their)potential) interactions)will)bring)about)changes)within)the)invaded)community)in)the)future.) ) )
) 5) References$ ) Allan,)B.)F.,)H.)P.)Dutra,)L.)S.)Goessling,)K.)Barnett,)J.)M.)Chase,)R.)J.)Marquis,)G.)Pang,)G.)A.) Storch,)R.)E.)Thach,)and)J.)L.)Orrock.)2010.)Invasive)honeysuckle)eradication)reduces) tick;borne)disease)risk)by)altering)host)dynamics.)Proceedings)of)the)National) Academy)of)Sciences)107:18523–18527.) Averill,)K.)M.,)D.)A.)Mortensen,)E.)A.)Smithwick,)and)E.)Post.)2016.)Deer)feeding)selectivity) for)invasive)plants.)Biological)Invasions:1–17.) Bartuszevige,)A.)M.,)D.)L.)Gorchov,)and)L.)Raab.)2006.)The)relative)importance)of)landscape) and)community)features)in)the)invasion)of)an)exotic)shrub)in)a)fragmented) landscape.)Ecography)29:213–222.) Bertness,)M.)D.,)and)R.)Callaway.)1994.)Positive)interactions)in)communities.)Trends)in) Ecology)&)Evolution)9:191–193.) Bloemer,)S.)R.,)E.)L.)Snoddy,)J.)C.)Cooney,)and)K.)Fairbanks.)1986.)Influence)of)deer)exclusion) on)populations)of)Lone)Star)ticks)and)American)dog)ticks)(Acari:)Ixodidae).)Journal) of)Economic)Entomology)79:679–683.) Castellano,)S.)M.,)and)D.)L.)Gorchov.)2013.)White;tailed)deer)(Odocoileus*virginianus)) disperse)seeds)of)the)invasive)shrub,)Amur)honeysuckle)(Lonicera*maackii).)Natural) Areas)Journal)33:78–80.) Christopher,)C.,)S.)Matter,)and)G.)Cameron.)2014.)Individual)and)interactive)effects)of)Amur) honeysuckle)(Lonicera*maackii))and)white;tailed)deer)(Odocoileus*virginianus))on) herbs)in)a)deciduous)forest)in)the)eastern)United)States.)Biological)Invasions) 16:2247–2261.) Cipollini,)K.,)E.)Ames,)and)D.)Cipollini.)2009.)Amur)honeysuckle)(Lonicera*maackii)) management)method)impacts)restoration)of)understory)plants)in)the)presence)of) white;tailed)deer)(Odocoileus*virginiana).)Invasive)Plant)Science)and)Management) 2:45–54.) Collier,)M.)H.,)J.)L.)Vankat,)and)M.)R.)Hughes.)2002.)Diminished)plant)richness)and) abundance)below)Lonicera*maackii,)an)invasive)shrub.)American)Midland)Naturalist) 147:60–71.)
) 6) Côté,)S.)D.,)T.)P.)Rooney,)J.;P.)Tremblay,)C.)Dussault,)and)D.)M.)Waller.)2004.)Ecological) impacts)of)deer)overabundance.)Annual)Review)of)Ecology,)Evolution,)and) Systematics)35:113–147.) Deering,)R.)H.,)and)J.)L.)Vankat.)1999.)Forest)colonization)and)developmental)growth)of)the) invasive)shrub)Lonicera)maackii.)The)American)Midland)Naturalist)141:43–50.) EDDMapS.)2015.)Amur)honeysuckle)(Lonicera*maackii));)EDDMapS)state)distribution.)Early) Detection)&)Distribution)Mapping)System.))The)University)of)Georgia);)Center)for) Invasive)Species)and)Ecosystem)Health.) http://www.eddmaps.org/distribution/usstate.cfm?sub=3040.))Accessed)June)1,) 2015.) Fagan,)M.)E.,)and)D.)R.)Peart.)2004.)Impact)of)the)invasive)shrub)glossy)buckthorn)(Rhamnus* frangula)L.))on)juvenile)recruitment)by)canopy)trees.)Forest)Ecology)and) Management)194:95–107.) Frappier,)B.,)R.)T.)Eckert,)and)T.)D.)Lee.)2004.)Experimental)removal)of)the)non;indigenous) shrub)Rhamnus*frangula*(Glossy)Buckthorn):)effects)on)native)herbs)and)woody) seedlings.)Northeastern)Naturalist)11:333–342.) Gallina,)S.,)and)H.)Lopez)Arevalo.)2008.)Odocoileus*virginianus.)The)IUCN)Red)List)of) Threatened)Species)2008:)e.T42394A10691422.) García,)D.,)and)J.)R.)Obeso.)2003.)Facilitation)by)herbivore;mediated)nurse)plants)in)a) threatened)tree,)Taxus*baccata:)local)effects)and)landscape)level)consistency.) Ecography)26:739–750.) Gómez;Aparicio,)L.,)R.)Zamora,)J.)Castro,)and)J.)A.)Hódar.)2008.)Facilitation)of)tree)saplings) by)nurse)plants:)Microhabitat)amelioration)or)protection)against)herbivores?) Journal)of)Vegetation)Science)19:161–172.) Gorchov,)D.)L.,)S.)M.)Castellano,)and)D.)A.)Noe.)2014.)Long;distance)dispersal)and)diffusion) in)the)invasion)of)Lonicera*maackii.)Invasive)Plant)Science)and)Management)7:464– 472.) Gorchov,)D.)L.,)and)D.)E.)Trisel.)2003.)Competitive)effects)of)the)invasive)shrub,)Lonicera* maackii)(Rupr.))Herder)(Caprifoliaceae),)on)the)growth)and)survival)of)native)tree) seedlings.)Plant)Ecology)166:13–24.)
) 7) Graff,)P.,)M.)R.)Aguiar,)and)E.)J.)Chaneton.)2007.)Shifts)in)positive)and)negative)plant) interactions)along)a)grazing)intensity)gradient.)Ecology)88:188–199.) Guiden,)P.,)D.)Gorchov,)C.)Nielsen,)and)E.)Schauber.)2015.)Seed)dispersal)of)an)invasive) shrub,)Amur)honeysuckle)(Lonicera*maackii),)by)white;tailed)deer)in)a)fragmented) agricultural;forest)matrix.)Plant)Ecology)216:939–950.) Hartman,)K.)M.,)and)B.)C.)McCarthy.)2008.)Changes)in)forest)structure)and)species) composition)following)invasion)by)a)non;indigenous)shrub,)Amur)honeysuckle) (Lonicera*maackii).)The)Journal)of)the)Torrey)Botanical)Society)135:245–259.) Hirth,)D.)H.)1977.)Social)behavior)of)white;tailed)deer)in)relation)to)habitat.)Wildlife) Monographs:3–55.) Horsley,)S.)B.,)S.)L.)Stout,)and)D.)S.)deCalesta.)2003.)White;tailed)deer)impact)on)the) vegetation)dynamics)of)a)northern)hardwood)forest.)Ecological)Applications)13:98– 118.) Keane,)R.)M.,)and)M.)J.)Crawley.)2002.)Exotic)plant)invasions)and)the)enemy)release) hypothesis.)Trends)in)Ecology)&)Evolution)17:164–170.) Krivtsov,)V.)2004.)Investigations)of)indirect)relationships)in)ecology)and)environmental) sciences:)a)review)and)the)implications)for)comparative)theoretical)ecosystem) analysis.)Lugano)aftermath)and)contemporary)advances)in)Ecological)Modelling:) Selected)materials)from)the)IEMSS)2002)conference)174:37–54.) Lieurance,)D.,)and)D.)Cipollini.)2012.)Damage)levels)from)arthropod)herbivores)on)Lonicera* maackii*suggest)enemy)release)in)its)introduced)range.)Biological)Invasions)14:863– 873.) Lieurance,)D.,)and)D.)Cipollini.)2013.)Exotic)Lonicera)species)both)escape)and)resist) specialist)and)generalist)herbivores)in)the)introduced)range)in)North)America.) Biological)invasions)15:1713–1724.) Luken,)J.)O.,)and)J.)W.)Thieret.)1996.)Amur)honeysuckle,)its)fall)from)grace.)BioScience) 46:18–24.) Mack,)R.)N.,)D.)Simberloff,)W.)Mark)Lonsdale,)H.)Evans,)M.)Clout,)and)F.)A.)Bazzaz.)2000.) Biotic)invasions:)causes,)epidemiology,)global)consequences,)and)control.)Ecological) Applications)10:689–710.)
) 8) McEwan,)R.)W.,)M.)K.)Birchfield,)A.)Schoergendorfer,)and)M.)A.)Arthur.)2009.)Leaf)phenology) and)freeze)tolerance)of)the)invasive)shrub)Amur)honeysuckle)and)potential)native) competitors.)The)Journal)of)the)Torrey)Botanical)Society)136:212–220.) McShea,)W.)J.)2012.)Ecology)and)management)of)white;tailed)deer)in)a)changing)world.) Annals)of)the)New)York)Academy)of)Sciences)1249:45–56.) Merriam,)R.)W.,)and)E.)Feil.)2002.)The)potential)impact)of)an)introduced)shrub)on)native) plant)diversity)and)forest)regeneration.)Biological)Invasions)4:369–373.) Miller,)J.)H.)2006.)Nonnative)invasive)plants)of)southern)forests.)USDA)Forest)Service/UNL) Faculty)Publications:103.) Orrock,)J.)L.,)H.)P.)Dutra,)R.)J.)Marquis,)and)N.)Barber.)2015.)Apparent)competition)and) native)consumers)exacerbate)the)strong)competitive)effect)of)an)exotic)plant)species.) Ecology)96:1052–1061.) Palmer,)W.)L.,)G.)M.)Kelly,)and)J.)L.)George.)1982.)Alfalfa)losses)to)white;tailed)deer.)Wildlife) Society)Bulletin)10:259–261.) Peck,)L.)J.,)and)J.)E.)Stahl.)1997.)Deer)management)techniques)employed)by)the)Columbus) and)Franklin)County)Park)District,)Ohio.)Wildlife)Society)Bulletin)(1973;2006)) 25:440–442.) Reichard,)S.)H.,)and)P.)White.)2001.)Horticulture)as)a)pathway)of)invasive)plant) introductions)in)the)United)States.)BioScience)51:103–113.) Rooney,)T.)2009.)High)white;tailed)deer)densities)benefit)graminoids)and)contribute)to) biotic)homogenization)of)forest)ground;layer)vegetation.)Plant)Ecology)202:103– 111.) Rooney,)T.)P.)2001.)Deer)impacts)on)forest)ecosystems:)a)North)American)perspective.) Forestry)74:201–208.) Rooney,)T.)P.,)and)D.)M.)Waller.)2003.)Direct)and)indirect)effects)of)white;tailed)deer)in) forest)ecosystems.)Forest)Ecology)and)Management)181:165–176.) Russell,)F.)L.,)D.)B.)Zippin,)and)N.)L.)Fowler.)2001.)Effects)of)white;tailed)deer)(Odocoileus) virginianus))on)plants,)plant)populations)and)communities:)a)review.)The)American) Midland)Naturalist)146:1–26.)
) 9) Smit,)C.,)C.)Vandenberghe,)J.)Ouden,)and)H.)Müller;Schärer.)2007.)Nurse)plants,)tree)saplings) and)grazing)pressure:)changes)in)facilitation)along)a)biotic)environmental)gradient.) Oecologia)152:265–273.) Sotala,)D.)J.,)and)C.)M.)Kirkpatrick.)1973.)Foods)of)white;tailed)deer,)Odocoileus)virginianus,) in)Martin)County,)Indiana.)American)Midland)Naturalist)89:281–286.) Strauss,)S.)Y.)1991.)Indirect)effects)in)community)ecology:)Their)definition,)study)and) importance.)Trends)in)Ecology)&)Evolution)6:206–210.) Trisel,)D.)E.,)and)D.)L.)Gorchov.)1994.)Regional)distribution,)ecological)impact,)and)leaf) phenology)of)the)invasive)shrub)Lonicera)maackii.)Bulletin)of)the)Ecological)Society) of)America)75:231.) Vandenberghe,)C.,)C.)Smit,)M.)Pohl,)A.)Buttler,)and)F.)Freléchoux.)2009.)Does)the)strength)of) facilitation)by)nurse)shrubs)depend)on)grazing)resistance)of)tree)saplings?)Basic)and) Applied)Ecology)10:427–436.) Webster,)C.)R.,)M.)A.)Jenkins,)and)S.)Jose.)2006.)Woody)invaders)and)the)challenges)they) pose)to)forest)ecosystems)in)the)eastern)United)States.)Journal)of)Forestry)104:366– 374.) White,)E.)M.,)J.)C.)Wilson,)and)A.)R.)Clarke.)2006.)Biotic)indirect)effects:)a)neglected)concept) in)invasion)biology.)Diversity)and)Distributions)12:443–455.) Woods,)K.)D.)1993.)Effects)of)invasion)by)Lonicera*tatarica)L.)on)herbs)and)tree)seedlings)in) four)New)England)forests.)American)Midland)Naturalist:62–74.) Wootton,)J.)T.)1994a.)The)nature)and)consequences)of)indirect)effects)in)ecological) communities.)Annual)Review)of)Ecology)and)Systematics:443–466.) Wootton,)J.)T.)1994b.)Predicting)direct)and)indirect)effects:)an)integrated)approach)using) experiments)and)path)analysis.)Ecology)75:151–165.) Wootton,)J.)T.)2002.)Indirect)effects)in)complex)ecosystems:)recent)progress)and)future) challenges.)Structuring)Factors)of)Shallow)Marine)Coastal)Communities,)Part)I) 48:157–172.) )
) 10) Chapter$2.$Effects$of$an$Invasive$Shrub,$Lonicera)maackii,$and$a$ Generalist$Herbivore,$WhiteEtailed$Deer,$on$Forest$Floor$Plant$ Community$Composition$
Introduction$ Indirect)effects)are)defined)as)“how)one)species)alters)the)effect)that)another)species) has)on)a)third”)(Strauss)1991).)Studies)of)plant);)animal)interactions)have)typically)focused) on)direct)effects)(Krivtsov)2004,)Strauss)and)Irwin)2004),)rather)than)indirect)effects,)as))) indirect)effects)are)more)difficult)to)observe,)predict,)and)quantify))(Krivtsov)2004,)White)et) al.)2006).)))Indirect)effects)between)plant)and)animal)species)are)fundamental)to) understanding)the)workings)of)complex)ecosystems)(Wootton)2002))and)may)be)important) for)understanding)invasions)(White)et)al.)2006).))While)invasive)plants)and)generalist) herbivores)have)each)been)shown)to)have)direct)negative)effects)on)plant)communities,) their)indirect)and)interactive)effects)have)received)far)less)focus.) With)the)ever;increasing)prevalence)of)global)travel)and)trade,)humans)have) purposefully)or)accidentally)introduced)many)species)of)plants)and)animals)to)areas)that) are)outside)of)their)native)ranges)(Mack)et)al.)2000,)Keane)and)Crawley)2002).))Once)these) organisms)are)introduced,)many)become)disruptive)and)detrimental)to)the)environments) they)invade)(Mack)et)al.)2000).))Invasive)plant)species)can)be)introduced)through) horticulture,)through)plantings)for)erosion)control,)or)though)contamination)of)crop)seeds,) among)other)reasons)(Reichard)and)White)2001).))The)effects)of)invasive)plant)species)on) other)plant)species)have)been)widely)documented,)and)generally)determined)to)be)negative) in)nature)(Mack)et)al.)2000,)Vilà)et)al.)2011),)while)impact)on)animal)species)have)been) more)mixed)(Hayes)and)Holzmueller)2012).) One)such)invasive)plant)species)is)Lonicera*maackii*(Rupr.))Herder)(Caprifoliaceae),) Amur)Honeysuckle,*a)large)shrub)that)is)native)to)northeast)Asia)(Luken)and)Thieret)1996),) and)is)one)of)several)bush)honeysuckle)species)that)are)invasive)in)the)eastern)United) States.)The)current)distribution)of)L.*maackii)spans)the)eastern)half)of)the)United)States;)L.* maackii)is)considered)invasive)and)regulated)in)eight)of)those)states)(EDDMapS)2015).)))
) 11) Lonicera*maackii)seeds)are)primarily)dispersed)by)birds)(Bartuszevige)et)al.)2006),)but)can) also)be)dispersed)by)white;tailed)deer)(Castellano)and)Gorchov)2013,)Guiden)et)al.)2015).)) After)L.*maackii)invades,)it)negatively)affects)forest)composition)in)several) ways.))Under)L.*maackii)shrubs,)there)is)lower)cover)of)tree)species)(Collier)et)al.)2002))and) tree)seedlings)experience)decreased)growth)and)survival)(Gorchov)and)Trisel)2003),)which) may)negatively)influence)future)forest)composition.))Tree)species)are)less)dense)and)differ) in)composition)in)both)the)seedling)and)sapling)layers)in)areas)of)L.*maackii*invasion) (Hartman)and)McCarthy)2008).)))Herb)species)are)less)abundant)in)areas)of)L.*maackii) invasion)(Hartman)and)McCarthy)2008))and)under)L.*maackii)shrubs)(Collier)et)al.)2002)) and)demonstrate)decreased)growth)and)reproduction)under)L.*maackii)shrubs)(Gould)and) Gorchov)2000,)Miller)and)Gorchov)2004).)The)composition)of)the)herb)community)differs) between)areas)with)and)without)L.*maackii*(Hartman)and)McCarthy)2008).***The)negative) effects)of)L.*maackii*on)herb)cover)beneath)its)canopy)extend)to)common)and)uncommon) spring)and)summer)herb)species)(Collier)et)al.)2002).)))Lonicera*maackii)also)negatively) affects)plant)species)richness)under)its)canopy)(Collier)et)al.)2002))of)both)native)and)exotic) plant)species)(Orrock)et)al.)2015).))Hartman)and)McCarthy)(2008))found)that)there)was)a) negative)effect)of)L.*maackii*on)tree,)but)not)herb,)species)richness)in)areas)of)invasion.) Additionally,)leaves)of)L.*maackii*decompose)more)rapidly)than)native)tree)species) (Poulette)and)Arthur)2012),)leading)to)decreased)litter)mass)and)increased)bare)ground,) factors)often)required)for)the)establishment)of)other)non;native)species)(Bartuszevige)et)al.) 2007,)Oswalt)and)Oswalt)2007,)Kuhman)et)al.)2013).) An)additional)factor)driving)changes)in)forest)ecosystems)in)the)United)States)and) Canada)is)high)density)of)white;tailed)deer,)Odocoileus*virginianus,*hereafter)referred)to)as) deer.))Deer)are)generalist)herbivores,)consuming)many)different)species)of)algae,)fungi,) herbs,)shrubs,)and)trees)(Atwood)1941,)Sotala)and)Kirkpatrick)1973).)Current)deer) densities)in)many)parts)of)the)United)States)are)well)above)historical)levels,)often)30;50) deer/km2)where)not)managed)(McShea)2012),)after)rebounding)from)widespread) population)declines)and)extirpations)in)the)early;to;mid)1800s)(Sotala)and)Kirkpatrick) 1973,)Côté)et)al.)2004).))This)overabundance)of)deer)has)negatively)impacted)tree)survival,) recruitment,)growth,)and)regeneration)(Russell)et)al.)2001,)Meiners)and)Martinkovic)2002,) Horsley)et)al.)2003,)Côté)et)al.)2004,)Hartman)and)McCarthy)2008,)Csigi)and)Holzmueller)
) 12) 2015))and)the)forestry)industry)through)browse)on)economically)important)tree)species) (Horsley)et)al.)2003,)Côté)et)al.)2004).))At)even)moderate)densities,)deer)greatly)alter) structure)and)composition)of)forests)(Rooney)2009).)At)densities)as)low)as)3;10)deer/km2) deer)negatively)impact)preferred)browse)species)(Horsley)et)al.)2003,)Côté)et)al.)2004).) These)preferences)for)forage)can)influence)the)winners)and)losers)of)competition)over)time,) acting)as)a)driver)of)change)(Wiegmann)and)Waller)2006),)and)increasing)the)prevalence)of) grasses,)ferns)and)other)species)deer)find)unpalatable)(Horsley)et)al.)2003,)Rooney)2009).)) Deer)have)also)been)shown)to)negatively)impact)plant)species)richness)(Wiegmann)and) Waller)2006,)Rooney)2009,)Relva)et)al.)2010,)Habeck)and)Schultz)2015).))Additionally,) areas)with)abundant)deer)show)increased)decomposition)rates)and)decreased)litter)mass) when)compared)to)areas)of)deer)exclosure)(Bressette)et)al.)2012).)) While)the)individual)direct)effects)of)deer)and)invasive)shrubs)on)forest) communities)are)well)studied,)only)a)modest)number)of)studies)have)investigated)indirect) effects)of)deer)on)invasive)plants,)or)interactive)effects)of)deer)and)invasive)plants)on)plant) communities.)Direct)effects)of)deer)on)invasive)plants)have)generally)been)investigated)in) the)form)of)consumption)of)the)invader)by)deer)(Schierenbeck)et)al.)1994).))Studies)of) indirect)effects)of)deer)on)invasive)plants)suggest)that)some)invasive)plants)are)facilitated,) even)obligatorily)so,)by)overabundant)deer)in)forest)settings)(Knight)et)al.)2009,)Kalisz)et)al.) 2014).))) Studies)of)simultaneous)and)interactive)effects)of)deer)and)invasive)plants)are)rare.))) Aronson)and)Handel)(2011))investigated)the)interactive)effects)of)deer)exclosure)and) removal)of)the)invasive)grass,)Microstegium*vimineum,*on)plant)communities,)finding) significant)effects)of)M.*vimineum)removal,)but)not)deer)or)their)interaction,)on)plant) species)richness.))Waller)and)Maas)(2013))investigated)the)effects)of)Alliaria*petiolata)and) deer)on)growth)and)survival)of)one)tree)species)and)four)herb)species.))Direct)negative) effects)of)deer)and)A.*petiolata)were)noted)on)survival)of)one)herb)species)and)growth)of)all) herb)species.)Negative)effects)of)deer)and)A.*petiolata)were)additive)for)one)herb)species) (Waller)and)Maas)2013).))There)was)a)significant)interaction)of)treatments)on)growth)of)the) tree)species,)with)greater)negative)effects)of)A.*petiolata)in)the)absence)of)deer)(Waller)and) Maas)2013).)* Additionally,)a)few)studies)have)suggested)interactive)effects)of)L.*maackii*and)deer)
) 13) on)native)plant)communities.)Christopher)et)al.)(2014))found)interactive)effects)of)deer)and) L.*maackii*on)the)abundance)of)annuals)and)spring)perennials:)where)L.*maackii)was) present)there)was)no)effect)of)deer)on)the)abundance)of)either)functional)group,)but)where) L.*maackii*was)removed)or)absent)deer)decreased)the)abundance)of)both.)There)was)also)an) interactive)effect)of)L.*maackii*and)deer)on)leaf)count)of)the)spring)perennial)herb)species,) Maianthemum*racemosum,)where)leaf)count)was)reduced)by)L.*maackii*when)deer)were) excluded,)but)there)was)no)reduction)of)leaf)count)when)deer)were)present.))Conversely,) Orrock)et)al.)(2015))found)no)significant)interactions)of)mammalian)consumers)(including) deer))and)L.*maackii*on)native)species)abundance)and)richness)or)exotic)species)abundance) or)richness.))However,)when)planted)Acer*saccharum)seedlings)were)accessible)to)deer,* their)biomass)was)much)greater)where*L.*maackii)shoots)were)present)than)where)they) had)been)removed;)where)seedlings)were)protected)from)deer,)their)mass)was)slightly) lower)where)L.*maackii*shoots)were)present)(Gorchov)and)Trisel)2003).))That)suggests)L.) maackii)protects)plants)from)deer)damage,)as)does)Cipollini)et)al.'s)(2009))finding)that)) herbaceous)plants)had)higher)reproduction)and)greater)size)beneath)both)standing)dead) and)living)shrubs)of)L.*maackii.**)Conversely,)Hartman)and)McCarthy)(2008))and) Christopher)et)al.)(2014))found)no)significant)effects)of)protection)from)deer)browse)on) tree)seedling)growth)and)survival.)) If)deer)and)L.*maackii)individually)negatively)affect)forest)floor)plants)(mechanisms) reviewed)above))without)interactive)effects)of)these)two)drivers,)these)effects)are)‘additive’) (Figure)1A).)I)predict)that)compositional)changes)should)follow)this)trend,)as)deer)and)L.* maackii)affect)composition)in)different)ways.))Specifically,)I)predict)species)unpalatable)to) deer)will)be)more)abundant)in)areas)with)deer)browse)than)in)areas)without)deer)(Horsley) et)al.)2003).))In)areas)with)L.*maackii*shrub)cover,)I)predict)a)large)negative)impact)on)tree) species)abundance,)and)a)smaller)negative)effect)on)forb)species)abundance)(Collier)et)al.) 2002,)Hartman)and)McCarthy)2008).))As)shading)by)the)early)leaf)flush)of)L.*maackii) hinders)spring)perennials’)access)to)sunlight)(Miller)and)Gorchov)2004),)I)also)predict) spring)perennials)will)be)more)impacted)by)L.*maackii*than)summer)perennials.) Alternatively,)deer)and)L.*maackii)may)impact)forest)plants)in)a)synergistic)manner.))) Synergistic)interactions)are)implied)if)the)joint)negative)effects)of)deer)and)L.*maackii*are) more)negative)than)the)sum)of)the)two)individual)effects)(Figure)1B).))Biologically,)this)can)
) 14) come)about)through)indirect)effects;)this)may)occur)if)deer)are)attracted)to)areas)with)L.* maackii*(refuge;mediated)apparent)competition,)sensu)Orrock)et)al.)(2010)),)if)plant) responses)to)shading)by)L.*maackii)makes)them)more)susceptible)to)deer)browse,)or)if)plant) responses)to)deer)browse)makes)them)less)tolerant)to)shading)by)L.*maackii.*** Antagonistic)interactions)occur)if)the)joint)negative)effects)of)deer)and)L.*maackii)are) less)negative)than)the)sum)of)the)two)individual)effects)(Figure)1C).))Biologically,)this)can) also)come)about)through)indirect)effects,)for)example,)if)deer)avoid)browsing)in)areas)with) L*maackii,)e.g.,)due)to)the)physical)barrier,)so)that)L.*maackii*decreases)the)negative)effect)of) deer)(Gorchov)and)Trisel)2003,)Cipollini)et)al.)2009),)or)if)deer)browse)on)L.*maackii,* reducing)its)cover)and)therefore)reducing)its)shading)of)other)plant)species.))I)predict)that) deer)and)L.*maackii*will)show)antagonistic)interactions)on)tree)seedlings)and)forest)floor) plant)species)richness,)with)deer)avoiding)areas)of)L.*maackii,*allowing)for)increased) survival)and)growth)of)tree)seedlings)and)other)preferred)species.)))
Methods$
Field)Methods) This)study)utilized)deer)exclosures)and)paired)deer)access)plots)arranged)in)a)split; plot)design)in)five)sites)in)the)Miami)University)Natural)Areas,)Butler)County,)Ohio)(39°)29’) ;)39°)31’)N,)84°)42’);)84°)43’)W,)Figure)2).))Sites)were)separated)by)≥)1)km)and)chosen)to) have)(1))closed;canopy)mature)deciduous)forest,)(2))level)topography,)and)(3))a)similar,) moderate)level)of)invasion)by)L.*maackii*(stem)basal)area)0.58)–)1.57)m2/ha,)Chapter)4).) Within)each)site,)two)similar)20m;by;20m)plots)were)selected;)one)each)was)randomly) assigned)to)deer)exclosure)and)deer)access)treatments.)Fencing)3m)high)was)placed)around) the)deer)exclosure)plots)in)summer)2010.)In)fall)of)2010,)basal)diameter)of)each)L.*maackii* stem)was)measured)in)each)half)(10)x)20)m))of)each)plot.))Lonicera*maackii)basal)areas)did) not)differ)between)treatment)types)(P)=)0.9305,)linear)mixed)effects)model,)with)site) included)as)a)random)factor,)Chapter)4).)Half)plots)were)randomly)assigned)to)be)L.*maackii) removed)or)intact)(Figure)).)Stems)of)L.*maackii)greater)than)1mm)in)diameter)were)clipped) at)the)base,)removed)from)the)plots,)and)stumps)treated)with)Tordon)RTU,)a)herbicide) composed)of)a)combination)of)5.4%)Picloram)(4;amino;3,)5,6;trichloropicolinic)acid))and) 20.9%)2,4;Dichlorophenoxyacetic)acid)(Dow)AgroScience)n.d.),)a)method)shown)to)be)the)
) 15) most)effective)in)killing)invasive)Rhamnus*cathartica)shrubs)(Pergams)and)Norton)2006).)) By)late)summer)2011,)some)of)the)treated)shrubs)had)re;sprouted;)these)new)stems)were) cut)and)treated)again)with)Tordon)RTU.)) I)sampled)the)forest)floor)vegetation)in)plots)twice)per)growing)season,)in)late) spring)and)mid)summer,)in)2011)–)2014)(Appendix)Table)A1).))Data)collected)in)2011)were) considered)the)baseline)for)comparisons)to)subsequent)years.))To)sample)the)vegetation)of) the)forest)floor,)a)modified)version)of)Daubenmire)plots)was)utilized)as)a)compromise) between)sampling)intensity)and)speed)of)sampling)(Abrahamson)et)al.)2011).)Each)half;plot) included)two,)20)m)transects)along)which)I)placed)nine)0.1m2)(20cm;by;50cm))subplots,) centered)every)2)m)(Appendix)Figure)A1).))Within)each)subplot,)all)species)<)1)m)tall)were) identified)and)the)area)of)each)was)estimated)using)an)acetate)sheet)with)sample)shapes)of) known)areas.)))Data)were)pooled)for)each)10)x)20)m)half;plot)and)were)used)to)calculate)1)) species)richness,)and)2))cover)of)each)species)under)1)m,)for)spring)and)summer)of)each) year)(Appendix)Table)A6).))In)every)year)and)season)after)spring)2012)I)also)determined) the)area)of)bare)ground)(visible)soil))for)each)sub;plot.))In)addition,)in)summer)2014,)the) number)of)reproductive)plants)of)Alliaria*petiolata,*an)invasive)biennial,)and)the)number)of) inflorescences)or)infructescences)of)Sanicula*odorata,*a)native)perennial,*were)censused)to) assess)effects)of)deer)and)L.*maackii*on)reproduction)of)these)species.)These)species)were) chosen)based)on)being)the)most)prevalent)invasive)(A.*petiolata))and)native)(S.*odorata)) species)in)the)plots)in)spring)of)2014.)))
Data)analysis) All)analyses)were)completed)using)R)version)3.1.1)(R)Core)Team)2014))utilizing)the) nlme)(linear)mixed)effects)models,)(Pinheiro)et)al.)2014),)vegan)(NMDS)and)PerMANOVA,) (Oksanen)et)al.)2008,)2015)),)BiodiversityR)(PerMANOVA,)(Kindt)and)Coe)2005),) indicspecies)(indicator)species)analysis,)(Caceres)and)Legendre)2009),)ggplot2)and)sciplot) packages)(Wickham)2009,)Morales)et)al.)2012).))Significance)was)determined)at)the)α))=) 0.05)level.$$) Species)richness)data)was)assessed)via)split;plot,)repeated)measures)analysis)of) variance)(ANOVA))utilizing)linear,)mixed)effects)models.))After)testing)for)normality)and) skewness,)richness)was)modeled)individually)by)season)as)spring)or)summer)Richness)=)
) 16) deer*L.*maackii*year)+)error,)with)deer)and)L.*maackii)treatments,)as)well)as)year,)as)fixed) effects,)and)site)as)a)random)effect.))If)the)3)way)interaction)was)non;significant,)that)term) was)then)dropped)from)the)model,)leaving)spring)or)summer)richness)=)deer*L.*maackii))+) deer*year)+)L.*maackii**year)+)error.))) Bare)ground)was)assessed,)for)each)year)and)season,)via)split;plot)ANOVA)utilizing)a) linear,)mixed)effect)model,))bare)ground)=)deer*L.*maackii)+)error,)with)deer)and)L.*maackii) treatments)as)fixed)effects,)and)site)as)a)random)effect.))If)the)2)way)interaction)was)non; significant)(all)years)and)seasons)except)spring)2012),)it)was)dropped)from)the)model,) leaving)Bare)Ground)=)Deer)+)L.*maackii)+)Error.) Composition)data)were)analyzed)with)non;metric)multi;dimensional)scaling) (NMDS),)a)form)of)ordination.)Bray;Curtis)dissimilarity)was)used)to)compute)a)distance) matrix,)and)data)were)stratified)by)site.)The)significance)of)treatment)effects)was) determined)using)3)dimensions,)selected)to)keep)stress)under)0.15,)and)modeled)using) with)main)effects)of)L.*maackii)and)deer,)with)site)as)a)random)effect.)The)inclusion)of) additional)dimensions)was)not)warranted)by)a)decrease)stress)in)the)model.))Significance) for)the)whole;plot)factor)(deer))was)determined)using)the)nested.npmanova)function)from) the)BiodiversityR)package)(Kindt)and)Coe)2005),)as)the)adonis)function)from)the)vegan) package)(Oksanen)et)al.)2015))tests)the)whole;plot)factor)(deer))with)the)incorrect)residual) error)term)for)a)split;plot)design)(Brewer)2013).) ) For)each)year)and)season)combination)that)had)a)significant)treatment)effect)on) composition,)I)carried)out)an)indicator)species)analysis.))Indicator)species)analysis)uses) data)on)species)presence)or)absence)as)well)as)cover)to)assess)strength)and)statistical) significance)of)the)relationship)between)species)occurrence/abundance)and)groups)of)sites) receiving)different)treatments)(Hill)et)al.)1975,)Bakker)2008,)De)Cáceres)and)Legendre) 2009,)De)Cáceres)et)al.)2010).))Indicator)values)(IV))are)calculated)with)higher)values) indicating)species)that)only)occur)in)that)group)and)lower)values)indicating)species)that) occur)across)all)groups)(Bakker)2008).)Tests)were)run)with)999)permutations.) ) I)further)investigated)composition)for)those)years)and)seasons)with)significant) treatment)effects)on)species)composition)by)analyzing)the)cover)of)plant)functional)types.)) Species)were)assigned)to)the)functional)types)used)by)Christopher)et)al.)(2014),)according) to)Gleason)and)Cronquist)(1991))and)Hochstedler)et)al.)(2007).))These)groups)were)vines,)
) 17) shrubs,)trees,)graminoids,)ferns,)spring)perennials,)summer)perennials,)annuals,)and) biennials.))Percent)cover)of)each)functional)type)was)tested)with)split;plot)ANOVA,)using)a) linear)mixed;effects)model:))Cover)=)Deer)Treatment)*L.*maackii*Treatment.)Deer)and)L.* maackii*treatments)were)included)as)fixed)effects,)with)site)included)as)a)random)effect.) Functional)types)that)were)not)present)in)≥)5)half;plots)were)excluded)from)analyses.))) ) Similarly,)I)analyzed)cover)of)invasive)plant)species)with)split;plot)ANOVAs,) modeled)with)a)linear)mixed)effects)model,)including)deer)and)L.*maackii)as)fixed)effects,) and)site)as)a)random)effect.))Invasive)status)for)the)lower)48)states)was)determined)using) the)United)States)Department)of)Agriculture)PLANTS)database)(USDA)2015).))Cover)of)L.* maackii)was)excluded)from)the)sum)of)the)cover)of)all)invasive)species.))The)non;L.*maackii) invasive)cover)was)tested)with)a)split;plot)ANOVA,)including)deer)and)L.*maackii* treatments)as)fixed)effects)and)site)as)a)random)effect.))If)the)2;way)interaction)was)not) significant,)it)was)dropped)from)the)model.)) Initial)(2011))cover)of)A.*petiolata*and)S.*odorata)were)assessed)via)split;plot) ANOVA,)modeled)with)a)linear)mixed)effects)model,)including)deer)and)L.*maackii)as)fixed) effects,)and)site)as)a)random)effect.)Because)the)2;way)interaction)was)non;significant,)it) was)dropped)from)the)model,)leaving)cover)=)deer)+)L.*maackii)+)error.))Reproductive)data) for)these)two)herbs,)number)of)individuals)with)flowers)or)fruits,)did)not)meet)assumptions) for)parametric)analysis,)so)were)instead)analyzed)with)Mann;Whitney)tests.)
Results$$
Species)Richness)
) Removing)Lonicera*maackii)had)a)positive)effect)on)spring)(F1,59)=)17.26,)P)=)0.0001,)
Table)1,)Figure)4A))and)summer)(F1,59)=)20.24,)P)<)0.0001,)Figure)4B))species)richness)over) the)course)of)the)study.)))Spring)species)richness)differed)among)years)(F3,59)=)7.19,)P)=) 0.0003,)Appendix)Figure)A3),)though)summer)species)richness)did)not.))There)was)a)trend) for)an)interaction)between)deer)exclosure)and)L.*maackii*removal)on)summer)plant)species) richness)(F1,59)=)3.76,)P))=))0.0573))where)in)areas)of)L.*maackii*presence,)there)was)a)less) negative,)or)smaller,)effect)of)deer)on)plant)species)richness)than)in)areas)where)L.*maackii) was)removed.)There)were)no)other)effects)of)deer)or)the)deer*L.*maackii)interaction)on) spring)or)summer)plant)species)richness.)
) 18) Species)Composition) ) Deer)exclosure)affected)spring)species)composition,)as)measured)by)NMDS,)in)2012)
(PERMANOVA)F5)=)2.35,)P)=)0.026),)2013)(F5)=)2.26,)P)=)0.015),)and)2014)(F5)=)2.10,)P)=) 0.003),)but)not)in)2011.))Neither)L.*maackii)removal)nor)the)deer*L.*maackii)interaction) affected)spring)species)composition)in)any)year)(Table)2).))Lonicera*maackii*removal) affected)summer)species)composition)in)2013)(Modeled)F1)=)1.96,)P)=)0.021),)but)not)in)any) other)year.)Deer)exclosure)and)the)interaction)between)L.*maackii*and)deer,)did)not)affect) summer)species)richness)in)any)year.))Site)affected)species)composition)in)every)year)and) season)except)summer)2011)(Table)2,)Appendix)Table)A2,)Appendix)Figure)A3).)
Indicator)Species) For)those)seasons)where)composition)was)affected)by)L.*maackii)or)deer)(above),)I) carried)out)indicator)species)analyses)(Table)3).)Deer)exclosure)was)indicated)by)the) presence)of)Mayapple)(Podophyllum*peltatum)*in)spring)2012)(Indicator)Value)(IV))=)0.835,) p)=)0.011))and)spring)2013)(IV)=)0.765,)p)=)0.037))and)by)Sugar)Maple)(Acer*saccharum))in) spring)2014,)IV)=)9.819,)p)=)0.024).)There)were)no)indicators)of)deer)access.)))) ) Removal)of)L.*maackii*was)indicated)by)the)presence)of)Virginia)creeper) (Parthenocissus*quinquefolia,)IV)=)0.878,)p)=)0.007),)Solomon’s)Seal)(Polygonatum*biflorum.* IV)=)0.707,)p)=)0.035),)and)Wild)licorice)(Galium*circaezans,)IV)=)0.691,)P)=)0.047))in) summer)2013.))The)presence)of)L.*maackii*was)indicated)by)the)presence)of)small)(<)1m) high)*L.*maackii*plants)(IV)=)0.876,)p)=)0.003).))
Cover)of)Functional)Types) Effects)of)deer)and)L.*maackii)treatments)on)cover)of)plant)functional)groups)were) investigated)only)for)those)seasons)where)composition)was)affected)by)at)least)one)of)these) treatments)(see)Species*Composition).))Analyses)are)reported)in)Appendix)Table)A4.)
Spring)2012,)2013,)and)2014) The)removal)of)L.)maackii)resulted)in)reduced)cover)of)shrubs)below)1)m)in)the) spring)of)2012)(F1,9)=)9.03,)P)=)0.0148,)Figure)14).))The)most)common)shrub)encountered) was)L.)maackii,)accounting)for)63)±)8)%)(Mean)±)SE))of)shrub)cover)under)1)m)across)all) treatments.)Lonicera)maackii)removal)increased)the)cover)of)annual)plants)(F1,9)=)7.19,))P)
) 19) =)0.0252))and)spring)perennials)(F1,9)=)6.90,)P)=)0.0275))in)spring)2013,)as)well)as)trees) (F1,9)=)5.57,)P)=)0.0426))and)graminoids)(F1,9)=10.45,)P)=)0.0103,)Figure)14))in)spring) 2014.)))There)were)no)other)effects)of)L.)maackii)in)any)other)year)or)season)for)any) functional)types.) Deer)exclosure)did)not)affect)any)plant)functional)types)in)any)year.)))However,)there) was)an)interaction)between)L.*maackii)and)deer)on)tree)cover)in)spring)2013)(F1,8)=14.22,))P)) =))0.0055).))The)highest)tree)cover)occurred)in)plots)without)L.*maackii*or)white;tailed)deer,) while)tree)cover)was)similar)among)areas)with)the)other)three)treatment)combinations) (Figure)6).)
Summer)2013) Lonicera)maackii)removal)increased)the)cover)of)vines)(F1,9)=)16.80,)P)=)0.0027))in)the) summer)of)2013)(Figure)14)..))Additionally,)there)was)an)interaction)of)L.)maackii)and)deer) on)cover)of)tree)species)(F1,8)=)5.97,)P)=)0.0404);)the)highest)tree)cover)occurred)without) L.)maackii)or)white;tailed)deer,)however,)there)were)no)differences)between)areas)with)and) without)deer)where)L.)maackii)was)intact)(Figure)7).))This)pattern)is)the)same)as)in)spring) 2013.)))
Bare)Ground) ) There)was)greater)area)of)bare)ground)(no)leaf)litter))in)areas)of)deer)access)in) spring)2014)and)summer)2012,)2013)and)2014.)In)Spring)2013,)there)was)an)effect)of)L.* maackii*removal)as)well)as)interaction)of)deer)and)L.*maackii,)where)the)greatest)amount)of) bare)ground)was)in)areas)of)L.*maackii*intact)and)deer)access,)with)other)areas)having) approximately)the)same)(Figure)8,)Table)4).)
Invasive)species) ) The)total)cover)of)non;L.*maackii)invasive)plant)species)was)not)affected)by)L.* maackii,*deer,)or)their)interaction)in)spring)or)summer)in)any)year)(Appendix)Table)A4).)
Herb)reproduction* All)reproduction)of)S.*odorata)in)summer)2014)(44)of)44)reproducing)individuals)) occurred)in)areas)of)deer)exclosure;)the)number)of)reproducing)individuals)differed) between)deer)treatments)(Mann;Whitney)Test,)W)=)25,)p)=)0.015))with)no)influence)of)L.*
) 20) maackii)treatment)(Fig.)9).)))Almost)all)reproduction)of)A.*petiolata)in)summer)2014)(7)of)8) reproducing)individuals))occurred)in)areas)of)deer)access)(Figure)9,)Mann;Whitney)Test,)W) =)71.5,)p)=)0.049).)))There)were)no)effects)of)L.*maackii,)deer,)or)their)interaction)on)S.* odorata*cover)in)summer)of)2011)or)A.*petiolata*cover)in)spring)2011,)suggesting)initial) equivalence)of)S.*odorata*and*A.*petiolata)cover)across)treatment)types)(Appendix)Table) A6).))
Discussion$ ) This)study)found)many)negative)direct)effects)of)L.*maackii)and)of)deer)on)plant) communities.))Interactive)effects)of)L.*maackii*and)deer)were)less)frequent,)but)in)each) instance,)provided)support)for)antagonistic)interactions.)))
Lonicera*maackii)direct)effects)) ) Removal)of)L.*maackii*had)a)positive)effect)on)plant)species)richness)during)the) course)of)the)study.))The)negative)impact)of)L.*maackii*on)species)richness*is)consistent) with)the)findings)of)Collier)et)al.)(2002).))) ) All)functional)groups)(trees,)annuals,)spring)perennials,)vines,)and)graminoids))had) greater)cover)in)areas)of)L.*maackii*removal,)with)the)exception)of)the)shrub)functional) type,)which)had)lower)abundance)in)areas)of)L.*maackii*removal.))These)effects)were)likely) responses)to)increased)light)availability)in)areas)of)L.*maackii*removal)(Luken)et)al.)1997,) Gorchov)and)Trisel)2003).)The)effects)of)L.*maackii*that)I)observed)on)tree)cover)were)both) as)predicted)and)consistent)with)the)findings)of)others)(Collier)et)al.)2002,)Gorchov)and) Trisel)2003,)Hartman)and)McCarthy)2008).))While)I)did)not)predict)the)significant)effect)of* L.*maackii*on)the)cover)of)graminoids,)this)finding)corroborates)the)trend)that)Christopher) et)al.)(2014))found)for)greater)cover)of)graminoids)in)areas)of)L.*maackii*removal.)I)also) found)that)spring,)but)not)summer,)perennials)had)less)cover)in)areas)with)L.*maackii*intact) compared)to)areas)of)L.*maackii*removed,)which)was)predicted)due)to)the)greater) sensitivity)of)spring)perennials)to)shading.)Miller)and)Gorchov)(2004))found)no)effect)of)L.* maackii*on)spring)perennial)survival,)though)they)did)find)greater)growth)and)reproduction) in)areas)of)L.*maackii*removal*while)Collier)et)al.)(2002))and)Christopher)et)al.)(2014)) found)L.*maackii*effects)on)abuncance)of)both)spring)and)summer)perennials.)))
) 21) ) The)effect)of)L.*maackii*on)the)area)of)bare)ground)was)only)significant)in)the)spring) of)2013,)even)though)L.*maackii)leaves)decompose)at)an)accelerated)rate)compared)to)other) native)species)(Poulette)and)Arthur)2012).))As)bare)ground)often)promotes)the) establishment)of)other)non;native)species)(Bartuszevige)et)al.)2007,)Oswalt)and)Oswalt) 2007,)Kuhman)et)al.)2013),)this)was)likely)a)contributing)factor)to)the)lack)of)effect)of)L.* maackii*on)the)cover)of)other)invasive)species.)))
Deer)Direct)Effects) ) Deer)exclosure)significantly)affected)plant)species)composition)in)every)year)after) the)first,)a)finding)similar)to)those)of)Horsley)et)al.)(2003))and)Rooney)(2009).))Indicators)of) areas)of)deer)exclosure)included)Podophyllum*peltatum*and)Acer*saccharum.**The)lower) cover)of)P.*peltatum)in)the)deer)access)plot)was)likely)due)to)trampling)by)deer,)since)P.* peltatum)was)often)found)knocked)over)and)is)unpalatable)to)deer)(Rawinski)and)Square) 2008,)Heckel)et)al.)2010).))A)study)with)another)unpalatable)herb,)Arisaema*triphyllum,) suggested)that)unpalatable)herbs)may)fail)to)benefit)from)high)deer)consumptive)pressure,) instead)decreasing)in)overall)prevalence,)possibly)due)to)increased)soil)compaction)in)areas) of)deer)access)(Heckel)et)al.)2010).))Acer*saccharum*seedling)density)has)been)previously) shown)to)increase)in)the)absence)of)deer)(Long)et)al.)2007).))) Area)of)bare)ground)was)greater)in)areas)of)deer)access)in)nearly)every)year)and) season)it)was)measured.)While)this)bare)ground)was)not)quantified)until)summer)2012,)the) effect)size)was)larger)in)2014)than)2012,)suggesting)an)increase)over)time)(Figure)8).) Similar)effects)have)been)reported)for)deer)on)bare)ground)(Knight)et)al.)2009))and)leaf) litter)depth)(Heckel)et)al.)2010),)and)are)likely)caused)by)increased)leaf)fragmentation)by) deer)trampling.) I)found)that)all)reproduction)of)S.*odorata)occurs)in)areas)of)deer)exclosure.))This)is) contrary)to)the)findings)of)Christopher)et)al.)(2014))who)found)that)there)was)no)effect)of) deer)exclosure)on)the)abundance)of)S.*odorata,)but)a)significant)effect)of)L.*maackii)removal.) However,)Knight)et)al.)(2009))found)that)much)smaller)proportion)of)native)plants)flower) in)areas)of)deer)access)compared)to)areas)of)deer)exclusion.))I)attribute)the)lack)of)S.* odorata)reproduction)in)areas)of)deer)access)to)deer)grazing)on)the)inflorescences)of)S.* odorata,*as)I)observed)stems)in)areas)of)deer)access)often)had)just)the)stem)tips)with)
) 22) inflorescences)removed.)Deer)florivory)can)be)extensive,)up)to)88;90%)of)all)floral) structures)in)one)species)of)Iris*experienced)florivory*(Geddes)and)Mopper)2006,)Wang)and) Mopper)2008),*and)can)negatively)effect)seed)production)and)resource)allocation)in) flowering)plants.)This)stark)dichotomy)of)plant)reproduction)may)eventually)lead)to)S.* odorata*only)occurring)in)and)near)areas)of)deer)exclosure,)though)established)individuals) may)persist)for)some)time.))))) I)found)the)opposite)trend)for)A.*petiolata.**All)reproducing)individuals)of)A.*petiolata) except)for)one)occurred)in)areas)of)deer)access.))This)is)in)congruence)with)findings)of) Kalisz)et)al.)(2014))that)deer)facilitate)growth)and)reproductive)success)of)A.*petiolata,)and) that)A.*petiolata)populations)declined)towards)local)extinction)over)six)years)of)deer) exclosure.)) I)did)not)find)any)effects)of)deer)exclosure)on)species)richness.))This)finding) contrasts)with)those)of)Wiegmann)and)Waller)(2006))and)Rooney)(2009),)who)found) decreased)richness)in)areas)of)deer)access.))Habeck)and)Schultz)(2015),)however,) completed)a)meta;analysis)of)25)studies)testing)the)effects)of)deer)on)understory)plant) communities,)and)found)no)significant)effect)of)deer)exclosure)on)plant)species)richness.)))
Interactions) There)was)an)interaction)between)L.*maackii*and)deer)on)tree)seedling)cover)in) spring)and)summer)of)2013.))The)direction)of)this)interaction)was)that)where)L.*maackii* was)intact,)there)were)no)differences)in)tree)seedling)cover)between)areas)of)deer) exclosure)and)access;)but)where)L.*maackii*was)removed,)there)was)greater)cover)of)tree) seedlings)in)areas)of)deer)exclosure)than)deer)access.))This)suggests)that)L.*maackii* mitigates)the)negative)effect)of)deer)browse)on)tree)seedling)cover,)and)facilitates)tree) seedling)survival)and)growth)(see)also)Chapter)3).))) There)was)also)a)trend)for)an)interaction)between)deer)and)L.*maackii*on)plant) species)richness)in)summer,)though)this)was)not)a)significant)effect)(F3,59)=)3.76,))P)=) 0.0573,)Table)1,)Appendix)Figure)A5).))In)areas)deer)exclosure,)there)was)much)lower) species)richness)in)areas)of)L.*maackii)intact)than)in)areas)of)L.*maackii*removal.))In)areas)of) deer)access,)species)richness)was)similar)between)L.*maackii)treatments.)This)pattern) suggests)deer)mitigate)the)negative)effect)of)L.*maackii.))A*possible)mechanism)for)this)
) 23) mitigation)is)that)deer)herbivory)on)L.*maackii*is)sufficient)to)reduce)its)cover.)Martinod) (unpubl.))found)that)nearly)5%)of)first;year)twigs)of)L.*maackii*in)this)study)area)showed) signs)of)deer)browse)between)bud)break)of)2015)and)May)2015.)I)found)that)where)deer) were)excluded,)there)was)higher)L.*maackii*leaf)frequency)at)heights)of)0.5)–)1.5)m)(see) Chapter)4),)within)the)0.3)–)2)m)height)range)in)which)deer)typically)browse)(Rooney)and) Waller)2003).))Additionally,)there)was)greater)L.*maackii)basal)area)growth)from)2010)to) 2015)in)deer)exclosures)(Chapter)4),)suggesting)that)deer)reduce)L.*maackii)growth)as)well) as)cover.)))
Support)for)Hypotheses) The)interaction)between)L.*maackii*and)deer)on)both)tree)cover)and)summer)plant) species)richness)was)antagonistic)in)nature)(Figure)1C),)with)areas)of)both)deer)and)L.* maackii*presence)having)a)less)negative)overall)effect)on)the)response)variables)than)would) be)predicted)from)the)additive)response)of)the)two)treatments)by)themselves.))This)pattern) could)be)due)to)either)lower)deer)browse)in)areas)with)L.*maackii,)due)to)the)physical) barrier,)so)that)L.*maackii*lessens)the)negative)effect)of)deer,)or)deer)browsing)on)L.* maackii,*reducing)its)cover)and)therefore)reducing)its)shading)of)other)plant)species,)so)that) deer)lessen)the)negative)effect)of)L.*maackii.))) The)mitigation)of)the)L.*maackii*effect)by)deer)is)supported)by)the)trend)of)an) interaction)between)L.*maackii*and)deer)on)summer)species)richness.))The)interactive) effects)of)L.*maackii*and)deer)on)tree)seedling)cover)support)mitigation)of)the)deer)effect)by) L.*maackii.**)Since)the)latter)response)was)statistically)significant,)I)posit)that)the)primary) interaction)between)deer)and)L.*maackii*is)the)mitigation)of)the)deer)effect)by)L.*maackii.** Previous)studies)have)suggested)that)interactions)between)L.*maackii)and)deer)may)confer) protection)from)browse)to)plants)under)the)canopy)of)L.*maackii.*Gorchov)and)Trisel) (2003),)while)testing)the)effects)of)L.*maackii*on)A.)saccharum*seedlings,)found)that)where) seedlings)were)accessible)to)deer,)A.*saccharum*mass)was)much)greater)where)L.*maackii) shoots)were)present.))Where)seedlings)were)fenced)to)protect)them)from)deer,)A.* saccharum)mass)was)only)slightly)higher)where)L.*maackii)shoots)were)absent.))Cipollini)et) al.)(2009))found)that)killing)L.*maackii)while)leaving)the)dead)stems)intact)led)to)less) reduction)of)seedlings)of)an)herb)species)by)deer)compared)to)areas)where)L.*maackii*was)
) 24) removed,)suggesting)that)standing)dead)shrubs)of)L.*maackii)provide)some)protection)from) deer)damage.))))) ) There)was)no)support)for)the)null)hypothesis)of)additive)effects)(Figure)1A))between) deer)and)L.*maackii.**There)were)no)instances)where)both)deer)and)L.*maackii)effects)were) significant)in)the)same)year)and)season.))There)also)was)no)evidence)from)this)study)to) support)synergistic)effects)between)L.*maackii)and)deer)(Figure)1B))as)the)combined)effects) were)never)more)negative)than)would)be)predicted)from)the)sum)of)the)two)effects.))) Our)results)suggest)that)considering)indirect)effects,)such)the)mitigation)of)the) negative)effect)of)deer)on)tree)seedlings*by)L.*maackii,)is)fundamental)to)understanding)the) roles)of)invasive)plants)and)over)abundant)herbivores)in)complex)natural)systems.)These) effects)are)rarely)simple,)and)could)drastically)impact)the)outcomes)of)management) initiatives.))For)example,)management)of)either)L.*maackii*or)deer)without)also)managing) the)other)factor)may)result)in)no)appreciable)change)in)the)plant)community,)particularly)in) terms)of)tree)recruitment)or)species)richness.))Our)study)shows)that)while)there)are) negative)impacts)of)both)herbivores)and)invasive)shrubs)on)native)plant)community) parameters,)the)combined)effects)are)less)negative)than)would)be)predicted)additively.))) ) )
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) 29) Oksanen,)J.,)F.)G.)Blanchet,)R.)Kindt,)P.)Legendre,)P.)R.)Minchin,)R.)B.)O’Hara,)G.)L.)Simpson,)P.) Solymos,)M.)H.)H.)Stevens,)and)H.)Wagner.)2015.)vegan:)community)ecology)package.) Oksanen,)J.,)Roeland)Kindt,)Pierre)Legendre,)Bob)O’Hara,)Gavin)L.)Simpson,)M.)Henry)H.) Stevens,)and)Helene)Wagner.)2008.)The)vegan)package:)community)ecology)package) in)R.) Orrock,)J.)L.,)H.)P.)Dutra,)R.)J.)Marquis,)and)N.)Barber.)2015.)Apparent)competition)and) native)consumers)exacerbate)the)strong)competitive)effect)of)an)exotic)plant)species.) Ecology)96:1052–1061.) Orrock,)J.)L.,)R.)D.)Holt,)and)M.)L.)Baskett.)2010.)Refuge;mediated)apparent)competition)in) plant–consumer)interactions.)Ecology)Letters)13:11–20.) Oswalt,)C.)M.,)and)S.)N.)Oswalt.)2007.)Winter)litter)disturbance)facilitates)the)spread)of)the) nonnative)invasive)grass)Microstegium*vimineum)(Trin.))A.)Camus.)Forest)ecology) and)management)249:199–203.) Pergams,)O.)R.)W.,)and)J.)E.)Norton.)2006.)Treating)a)single)stem)can)kill)the)whole)shrub:)a) scientific)assessment)of)buckthorn)control)methods.)Natural)Areas)Journal)26:300– 309.) Pinheiro,)J.,)D.)Bates,)S.)DebRoy,)D.)Sarkar,)and)R)Core)Team.)2014.)nlme:)linear)and) nonlinear)mixed)effects)models.) Poulette,)M.)M.,)and)M.)A.)Arthur.)2012.)The)impact)of)the)invasive)shrub)Lonicera*maackii) on)the)decomposition)dynamics)of)a)native)plant)community.)Ecological) Applications)22:412–424.) Rawinski,)T.)J.,)and)N.)Square.)2008.)Impacts)of)white;tailed)deer)overabundance)in)forest) ecosystems:)an)overview.)USDA)Forest)Service,)Newton)Square,)PA.)Available)online) at:)http://www.)na.)fs.)fed.)us/fhp/special_interests/white_tailed_deer.)pdf.) R)Core)Team.)2014.)R:)a)language)and)environment)for)statistical)computing.)R)Foundation) for)Statistical)Computing,)Vienna,)Austria.) Reichard,)S.)H.,)and)P.)White.)2001.)Horticulture)as)a)pathway)of)invasive)plant) introductions)in)the)United)States.)BioScience)51:103–113.) Relva,)M.,)M.)Nuñez,)and)D.)Simberloff.)2010.)Introduced)deer)reduce)native)plant)cover)and) facilitate)invasion)of)non;native)tree)species:)evidence)for)invasional)meltdown.) Biological)Invasions)12:303–311.)
) 30) Rooney,)T.)2009.)High)white;tailed)deer)densities)benefit)graminoids)and)contribute)to) biotic)homogenization)of)forest)ground;layer)vegetation.)Plant)Ecology)202:103– 111.) Rooney,)T.)P.,)and)D.)M.)Waller.)2003.)Direct)and)indirect)effects)of)white;tailed)deer)in) forest)ecosystems.)Forest)Ecology)and)Management)181:165–176.) Russell,)F.)L.,)D.)B.)Zippin,)and)N.)L.)Fowler.)2001.)Effects)of)white;tailed)deer)(Odocoileus* virginianus))on)plants,)plant)populations)and)communities:)a)review.)The)American) Midland)Naturalist)146:1–26.) Sotala,)D.)J.,)and)C.)M.)Kirkpatrick.)1973.)Foods)of)white;tailed)deer,)Odocoileus*virginianus,) in)Martin)County,)Indiana.)American)Midland)Naturalist)89:281–286.) Strauss,)S.)Y.)1991.)Indirect)effects)in)community)ecology:)Their)definition,)study)and) importance.)Trends)in)Ecology)&)Evolution)6:206–210.) Strauss,)S.)Y.,)and)R.)E.)Irwin.)2004.)Ecological)and)evolutionary)consequences)of) multispecies)plant;animal)interactions.)Annual)Review)of)Ecology,)Evolution,)and) Systematics)35:435–466.) USDA.)2015,)August)5.)Welcome)to)the)PLANTS)database)|)USDA)PLANTS.) http://plants.usda.gov/java/.) Vilà,)M.,)J.)L.)Espinar,)M.)Hejda,)P.)E.)Hulme,)V.)Jarošík,)J.)L.)Maron,)J.)Pergl,)U.)Schaffner,)Y.) Sun,)and)P.)Pyšek.)2011.)Ecological)impacts)of)invasive)alien)plants:)a)meta;analysis) of)their)effects)on)species,)communities)and)ecosystems.)Ecology)letters)14:702– 708.) Waller,)D.)M.,)and)L.)I.)Maas.)2013.)Do)white;tailed)deer)and)the)exotic)plant)garlic)mustard) interact)to)affect)the)growth)and)persistence)of)native)forest)plants?)Forest)Ecology) and)Management)304:296–302.) Wang,)H.)G.,)and)S.)Mopper.)2008.)Separate)and)interacting)effects)of)deer)florivory)and) salinity)stress)on)Iris)herbivores.)Oikos)117:564–570.) White,)E.)M.,)J.)C.)Wilson,)and)A.)R.)Clarke.)2006.)Biotic)indirect)effects:)a)neglected)concept) in)invasion)biology.)Diversity)and)Distributions)12:443–455.) Wickham,)H.)2009.)ggplot2:)elegant)graphics)for)data)analysis.)Springer)New)York.)
) 31) Wiegmann,)S.)M.,)and)D.)M.)Waller.)2006.)Fifty)years)of)change)in)northern)upland)forest) understories:)Identity)and)traits)of)“winner”)and)“loser”)plant)species.)Biological) Conservation)129:109–123.) Wootton,)J.)T.)2002.)Indirect)effects)in)complex)ecosystems:)recent)progress)and)future) challenges.)Structuring)Factors)of)Shallow)Marine)Coastal)Communities,)Part)I) 48:157–172.) ) )
) 32) Table$1.)Repeated)measures)ANOVA)table)for)plant)species)richness.))Significant)treatment) effects)are)bolded.))Deer)refers)to)effect)of)deer)exclosure;)L.*maackii*refers)to)removal)of) Lonicera*maackii.)) ) Season) Treatment) df* den)df) F) P) Spring) )Deer) 1** 4)) 0.98686) ) 0.3767) ) ) Honeysuckle$ 1) 59$ 17.25998$ 0.0001$ ) Year$ 3) 59$ 7.18871$ 0.0003$ ) Deer:Honeysuckle) 1* 59) 2.85339) 0.0965) ) Deer:Year) 3* 59) 0.79133) 0.5036) ) Honeysuckle:Year) 3* 59) 1.49166) 0.2261) )Summer) )Deer) 1** 4)) 0.56674) ) 0.4934) ) ) Honeysuckle$ 1) 59$ 20.24474$ <.0001$ ) Year) 3* 59) 1.82478) 0.1525) ) Deer:Honeysuckle) 1* 59) 3.75884) 0.0573) ) Deer:Year) 3* 59) 1.48935) 0.2267) )) Honeysuckle:Year) 3* 59) 0.45095) 0.7176)
) )
) 33) Table$2.)Significance)of)treatment)effects)on)species)composition)of)forest)floor)plants)by) year)and)season.)))Values)determined)using)permutational)multivariate)analysis)of)variance) (Appendix)Table)A2).))Non;significant)p;values)are)reported)as)n.s.)
) 2011) 2012) 2013) 2014)
) Spring) Summer) Spring) Summer) Spring) Summer) Spring) Summer)
Deer) n.s.) n.s.) 0.026) n.s.) 0.015) n.s.) 0.003) n.s.)
L.*maackii* n.s.) n.s.) n.s.) n.s.) n.s.) 0.021) n.s.) n.s.)
Interaction) n.s.) n.s.) n.s.) n.s.) n.s.) n.s.) n.s.) n.s.)
Site) 0.029) n.s.) 0.017) 0.014) 0.039) 0.002) 0.023) 0.001)
)
) 34) Table&3.!!Indicator!species!of!significant!compositional!treatment!effects.!!!
Indicator! Common!! Year! Season! Treatment! Indicator!of:! Value! P! Name! Scientific!Name! 2012! Spring! Deer! ! Exclosure! ! 0.835! ! 0.011! ! Mayapple! ! Podophyllum*peltatum! * ! 2013! Spring! ! Deer! ! Exclosure! ! 0.765! ! 0.037! ! Ma! yapple! Podophyllum*peltatum* * ! 2014! Spring! ! Deer! ! Exclosure! ! 0.819! ! 0.024! ! Sugar!Maple! ! Acer*saccharum* * ! 2013! ! Summer! ! ! ! ! ! ! * ! !Honeysuckle! !Intact! 0.876! ! 0.003! ! !Amur! *Lonicera*maackii* Honeysuckle! ! ! ! Removed! 0.878! 0.007! Virginia!Creeper! Parthenocissus*quinquefolia* ! ! Removed! 0.707! 0.035! Solomon's!Seal! Polygonatum*biflorum* !! !! !! Removed! 0.691! 0.047! Wild!Licorice! Galium*circaezans* !
!
!
!
! 35! Table&4.!Significance!of!treatment!effects!on!area!of!bare!ground!by!year!and!season.!!! Values!determined!using!split9plot!analysis!of!variance.!!&
Season! Year! !! numDF! denDF! F9value! p9value! Summer! 2012! Deer! & 1!& 4!& 8.591542! & 0.0428! & & & Honeysuckle! 1! 9! 3.056225! 0.1144! Spring! ! ! 2013! Deer! ! 1!! 4!! 4.448971! ! 0.1026! ! & & Honeysuckle& 1& 8& 6.339356& 0.0359& ! ! Deer:Honeysuckle& 1& 8& 11.985345& 0.0085& Summer! ! ! 2013! Deer! & 1!& 4!& 10.326272! & 0.0325! & & & Honeysuckle! 1! 9! 3.410826! 0.0978! Spring! ! ! 2014! Deer! & 1!& 4!& 8.272542! & 0.0452! & & & Honeysuckle! 1! 9! 1.500559! 0.2517! Summer! ! ! 2014! Deer! & 1!& 4!& 8.272542! & 0.0452! & !&! !&! Honeysuckle! 1! 9! 1.500559! 0.2517! !
!
! &
! 36! &
&
&
&
&
&
&
! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! Figure&1.!!Predictions!of!plant!community!responses.!!A.!Predictions!of!plant!community! responses!based!on!the!null!hypothesis!that!effects!of!deer!and!Lonicera)maackii!are! additive.!B.!Predictions!of!plant!community!responses!based!on!the!alternative!hypothesis! that!effects!of!deer!and!L.)maackii!are!non9additive!and!synergistic!in!nature.)C.!Predictions! of!plant!community!on!the!alternative!hypothesis!that!effects!of!deer!and!L.)maackii!are! non9additive!and!antagonistic!in!nature.!!
!
! 37! ! Figure&2.!!Map!of!study!sites!in!the!Miami!University!Natural!Areas.!!Pink!squares!denote! areas!of!deer!access!plots,!while!yellow!squares!denote!areas!of!deer!exclosure,!for!a!total! of!10!paired!plots!across!five!Natural!Areas.!!Natural!Areas!are!annotated!with!their!names.&! !
! 38! ! Figure&3.!!Schematic!of!study!design.!!Solid!lines!indicate!the!20!x!20!m!fenced!exclosure! area,!also!indicated!by!the!figure!of!the!deer!covered!by!the!no!symbol.!!Dotted!lines! indicate!completely!permeable!boundaries.!!The!shrub!covered!by!the!no!symbol!indicates! removal!of!L.)maackii.!!!
&
! 39! A)# Spring Species Richness B)# Summer Species Richness 2011 2012 2013 2014 2011 2012 2013 2014
25 Honeysuckle RE
25 PR
20
20
15 Richness Richness
15
10
10 Honeysuckle RE PR
AC EX AC EX AC EX AC EX AC EX AC EX AC EX AC EX Deer Deer ! Figure!4.!!Boxplots!of!A)!spring!and!B)!summer!plant!species!richness!across!the!4!years!of!the!study.!!Gray!boxes!represent! areas!with!Lonicera)maackii!intact,!white!boxes!represent!areas!with!L.)maackii)removed.!!Deer!treatment!is!shown!on!the!x? axis,!with!areas!with!deer!access!marked!as!AC,!areas!that!have!deer!exclosures!marked!with!EX.!
! 40! Spring 2012 Spring 2013 Annual Graminoid Shrub Spring Summer Tree Vine Annual Graminoid Shrub Spring Summer Tree Vine 30 Honeysuckle Honeysuckle 40 RE RE 30 PR 20 PR
20 10
PercentCover 10 PercentCover
0 0 AC EX AC EX AC EX AC EX AC EX AC EX AC EX AC EX AC EX AC EX AC EX AC EX AC EX AC EX Deer Deer
Spring 2014 Summer 2013 Biennial Graminoid Shrub Spring Summer Tree Vine Graminoid Shrub Summer Tree Vine 40 30 Honeysuckle Honeysuckle RE RE 30 PR PR 20 20
10
PercentCover PercentCover 10
0 0 CO EX CO EX CO EX CO EX CO EX CO EX CO EX AC EX AC EX AC EX AC EX AC EX Deer Deer ! Figure!5.!!Boxplots!of!cover!of!plant!functional!types!by!treatment!in!spring!2012,!2013,!and!2014,!and!summer!2013.!!Gray! boxes!represent!areas!with!Lonicera)maackii!intact,!white!boxes!represent!areas!with!L.)maackii!Removed.!!Black!circles!are! outliers.!!!Deer!treatment,!is!shown!on!the!x?axis,!with!areas!with!deer!access!marked!as!AC,!areas!that!have!deer!exclosures! marked!with!EX.!!!Functional!types!‘Spring’!and!‘Summer’!refer!to!spring!and!summer!perennials.!!! !
! 41! !
Tree Cover Spring 2013
Deer
EX
4 CO 3 Cover (Percent) Cover 2 1
PR RE
Honeysuckle Treatment !! Figure'6.!!Percent!tree!seedling!cover!in!Spring!2013.!!Interaction!plot!shows!mean!percent! tree!cover,!+>!SD,!by!treatment!type.!!Open!circles!represent!areas!of!deer!exclosure.!!Filled! circles!represent!areas!of!deer!access.!Lonicera)maackii!treatment,!is!shown!on!the!x>axis,! with!areas!with!L.)maackii!removed!represented!as!RE!and!areas!with!L.)maackii)present! with!PR,.!' ! '
! 42! !
Tree Cover Summer 2013 5 Deer
EX AC 4 3 Cover (Percent) Cover 2 1
PR RE
Honeysuckle Treatment !! Figure'7.!!Percent!tree!seedling!cover!in!Summer!2013.!!Interaction!plot!shows!mean! percent!tree!cover,!+>!SD,!by!treatment!type.!!Open!circles!represent!areas!of!deer! exclosure.!!Filled!circles!represent!areas!of!deer!access.!Lonicera)maackii!treatment,!is! shown!on!the!x>axis,!with!areas!with!L.)maackii!removed!represented!as!RE!and!areas!with! L.)maackii)present!with!PR,.!'
'
! !
! 43! !
Bare Ground Spring13 Spring14 Summer12 Summer13 Summer14 20
15
Honeysuckle 10 RE PR BareGroundPercent
5
0
AC EX AC EX AC EX AC EX AC EX Deer ! ! Figure'8.''Boxplots!of!percent!bare!ground!in!spring!2013!and!2014!and!summer!2012,! 2013,!and!2014.!!White!boxes!represent!areas!with!L.)maackii!removed;!gray!boxes! represent!areas!with!L.)maackii!present.!!Deer!treatment!is!shown!on!the!x>axis,!with!areas! with!deer!access!marked!as!AC!areas!that!have!deer!exclosures!marked!with!EX.!!
! !
! 44! !
Sanicula odorata Alliaria petiolata 10.0 Honeysuckle Honeysuckle PR PR RE RE
7.5 1.0
5.0
0.5
Number of Reproducing Individuals of Reproducing Number 2.5 Individuals of Reproducing Number
0.0 0.0
AC EX AC EX Deer Deer ! Figure'9.''Mean!number!(±SE)!of!reproductive!individuals!of'Sanicula)odorata!and!Alliaria) petiolata)per!plot!by!treatment!type.!Light!gray!bars!represent!areas!with!L.)maackii!intact;! Dark!Gray!bars!represent!areas!with!L.)maackii!Removed.!!Deer!treatment!is!shown!on!the! x>axis,!with!areas!with!deer!access!marked!as!AC,!areas!that!have!deer!exclosures!marked! with!EX.!!There!were!no!reproducing!individuals!of!S.)odorata!in!areas!with!deer!access,! and!there!was!only!one!reproducing!individual!of!A.)petiolata!in!areas!of!deer!exclosure.!!
' !
! 45! !
Chapter'3.'Are'native'tree'seedlings'facilitated'by'an'invasive'shrub' where'white>tailed'deer'are'abundant?'
Introduction' The!competitive,!negative!effects!of!various!invasive!shrub!species!on!tree!seedlings! are!well!known!(Woods!1993,!Merriam!and!Feil!2002,!Fagan!and!Peart!2004,!Frappier!et!al.! 2004,!Webster!et!al.!2006).!!However,!many!shrub!species!are!known!to!protect!tree! seedlings!in!areas!of!high!herbivore!presence!(Bakker!et!al.!2004,!Smit!et!al.!2007,!Jensen!et! al.!2012,!Perea!and!Gil!2014).!This!latter!interaction!is!an!example!of!facilitation,!a!positive! interaction!that!is!not!consumptive!in!nature!and!that!benefits!at!least!one!of!the!species! involved!(Merriam!and!Feil!2002).!!Plants!can!positively!interact!with!each!other!through! the!modification!of!physical!or!biotic!conditions,!which!can!include!the!facilitation!of!their! neighbors!through!protection!from!herbivore!browse!(Callaway!1995).!!It!has!more! recently!been!shown!that!positive!plant>to>plant!interactions,!previously!viewed!as! features!only!of!desert!environments,!are!important!in!areas!with!environmental! conditions!harsh!in!ways!other!than!extreme!water!limitation!(Bertness!and!Callaway! 1994).!Negative!interactions,!which!have!received!the!most!study!(Berkowitz!et!al.!1995,! Smit!et!al.!2007),!are!predicted!at!intermediate!levels!of!abiotic!stress,!while!facilitation!is! predicted!in!environments!that!are!more!stressful!to!plants!(Bertness!and!Callaway!1994).!!! Intense!herbivore!browse!pressure!is!a!stressor!that!can!prompt!facilitative!plant> to>plant!interactions!(Bertness!and!Callaway!1994,!Gómez>Aparicio!et!al.!2008).!In!a! forested!area!browsed!by!five!different!species!of!ungulate!herbivores,!García!and!Obeso! (2003)!found!that!the!threatened!yew!species!Taxus)baccata!L.!establishes!only!under!the! well!defended!shrub/small!tree,!Ilex)aquifolium!L.;!T.)baccatta)experienced!higher!seedling! survival!and!lower!herbivory!on!saplings!beneath!I.)aquifolium.!Jensen!et!al.!(2012)!found! reduced!browse!on!Quercus)robur)and!Q.)petraea!under!a!canopy!of!multiple!shrub!species,! including!structurally!protected!Rubus)idaeus,!in!areas!with!two!large,!ungulate!herbivores.!!! A!trend!that!emerges!is!that!well!defended!species!or!those!otherwise!not!preferred!as!a! food!source!facilitate!the!growth!of!their!neighbors!under!grazing!or!browsing!pressure! (Graff!et!al.!2007,!Smit!et!al.!2007,!Vandenberghe!et!al.!2009).!Therefore,!one!might!expect!
! 46! ! that!if!a!species!is!not!preferred!as!food!by!an!herbivore,!it!potentially!provides!a!refuge! from!herbivory!for!other!species,!facilitating!their!growth!and!survival.!! ! Deer!densities!in!many!parts!of!the!United!States!are!well!above!historical!levels,! often!30>50!deer/km2!where!not!managed!(McShea!2012).!!The!range!of!Odocoileus) virginianus!Zimmerman,!white>tailed!deer,!extends!into!each!of!the!contiguous!48!United! States,!Canada,!Mexico,!Central!America,!and!Northern!South!America!(Hirth!1977,!Russell! et!al.!2001).!!Estimates!of!pre>settlement!deer!densities!range!from!2>4!deer/km2!across! their!entire!North!American!range!(Rooney!2001),!and!have!increased!to!average!densities! of!approximately!10!deer!deer/km2!(Rooney!and!Waller!2003).!!! Deer!greatly!impact!the!areas!in!which!they!occur,!changing!structure!and! composition!of!forests!through!browse!on!important!tree!species!(Horsley!et!al.!2003,!Côté! et!al.!2004,!Rooney!2009).!!At!densities!as!low!as!3>10!deer!deer/km2,!deer!negatively! impact!preferred!browse!species!(Horsley!et!al.!2003,!Côté!et!al.!2004).!Their!preferences! for!forage!drive!changes!in!species!abundances,!decreasing!the!frequency!of!favored! species,!and!increasing!the!relative!prevalence!of!species!deer!find!unpalatable!(Horsley!et! al.!2003,!Wiegmann!and!Waller!2006,!Rooney!2009).!! ! One!species!of!shrub!that!potentially!functions!as!a!facilitator!species!is!Lonicera) maackii!(Rupr.)!Herder!(Caprifoliaceae),!Amur!Honeysuckle.!!!Lonicera)maackii!is!a!large! shrub!with!dense!foliage!that!grows!to!heights!of!up!to!6!meters!(Luken!and!Thieret!1996),! and!is!one!of!several!bush!honeysuckle!species!that!are!invasive!in!the!eastern!United! States!(Miller!2006).!Its!native!range!includes!China,!Japan,!Korea!and!Southeastern!Russia! (Luken!and!Thieret!1996).!!The!current!distribution!of!L.)maackii!spans!the!eastern!half!of! the!United!States;!L.)maackii!is!considered!invasive!and!regulated!in!8!of!those!states! (EDDMapS!2015).!While!the!browse!preference!by!deer!of!L.)maackii)is!not!well!studied,!its! leaves!have!low!palatability!for!insects!(Lieurance!and!Cipollini!2012,!2013),!and!the! closely!related!L.)morrowii!A.!Gray,)also!invasive,!)is!of!moderate!preference!to!deer!(Averill! et!al.!2016).!!Lonicera)maackii!has!many!negative!direct!on!plant!communities,!and!has! been!shown!to!decreases!growth,!survival,!density!and!cover!of!tree!seedlings!and!saplings! (Collier!et!al.!2002,!Gorchov!and!Trisel!2003,!Hartman!and!McCarthy!2008).!) ! Previous!studies!have!suggested!that!interactions!between!L.)maackii)and!deer!may! confer!protection!from!browse!to!plants!under!the!canopy!of!L.)maackii.)!Cipollini!et!al.!
! 47! !
(2009)!suggested!that!standing!dead!L.)maackii!shrubs!may!provide!protection!to!native! herbs!from!deer.!!This!structural!barrier!may!prevent!them!from!being!browsed.!Gorchov! and!Trisel!(2003),!while!testing!the!effects!of!L.)maackii)on!ACSA)seedlings,!found!that! where!seedlings!were!accessible!to!deer,!ACSA)mass!was!much!greater!where!L.)maackii! shoots!were!present.!!This!finding!suggests!that!L.)maackii!may!prevent!mass!loss!to!deer! herbivory.!!! ! I!investigated!whether!L.)maackii!provides!a!refuge!for!native!tree!species!from!deer! browse,!allowing!tree!seedlings!to!attain!greater!height!and!survival!when!planted!under!L.) maackii!shrubs!than!when!planted!away.!!!I!hypothesized!that!facilitation,!rather!than! competition,!is!the!dominant!interaction!between!L.)maackii!and!native!tree!species,!and! that!honeysuckle!provides!protection!to!tree!seedlings!from!heavy!deer!browse!(Figure!1).!!
Materials'and'Methods'
Field!Methods) In!order!to!test!whether!L.)maackii)provides!a!refuge!from!browsing,!I!measured!the! growth!and!survival!of!tree!seedlings!planted!under!individual!L.)maackii!shrubs,!with!and! without!deer!herbivory!(Figure!2).!!Within!the!Miami!University!Natural!Areas,!in!Oxford,! Ohio!(Butler!County),!USA,!I!selected!two!study!areas,!one!with!high!deer!density!(Western! Woods,!12.1!deer/km2!(Summer)!and!23.8!deer/km2!(Winter),!Barrett!2014),!and!one!with! lower!deer!density!(Kramer!Preserve,!9.9!deer/km2!(Summer)!and!6.7!deer/km2!(Winter),! Barrett!2014).!!In!each!Natural!Area,!10!large,!focal!shrubs!were!chosen!that!A)!were!>!2m! tall,!B)!had!>!1!stem,!C)!had!a!canopy!that!extended!down!to!!1!m!away!from!the!closest!tree,! and!F)!were!>!4!m!from!a!trail!(Figure!2).!!!Other!small!L.)maackii)plants!within!the!canopy! area!of!the!focal!shrub!were!removed!and!the!stems!were!treated!with!Tordon!RTU,!a! herbicide!composed!of!a!combination!of!5.4%!Picloram!(4>amino>3,5,6>trichloropicolinic! acid)!and!20.9%!2,4>Dichlorophenoxyacetic!acid!(Dow!AgroScience!n.d.),!a!method!shown! to!be!the!most!effective!method!to!kill!and!prevent!re>growth!of!invasive!shrub!Rhamnus) cathartica!(Pergams!and!Norton!2006).!!After!selection,!focal!shrubs!were!randomly! assigned!to!be!fenced!deer!exclosure!plots,!or!unfenced!deer!access!plots.!Deer!exclosure!
! 48! ! plots!were!surrounded!with!1.25!m!tall!black!plastic!mesh!fencing,!approximately!1!m!from! the!edge!of!the!shrub’s!canopy.!!Exclosures!were!established!May!5!–!23,!2014.!! !Bare!root!tree!seedlings!of!four!species!were!selected;!two!of!very!high!deer! preference!(Quercus!alba!L.!and!Carya)ovata!(Mill.)!K.Koch,!(Strole!and!Anderson!1992,! Wakeland!and!Swihart!2009)),!hereafter!QUAL!and!CAOV,!respectively,!one!of!moderate! deer!preference!(Acer)saccharum!Marshall,!(Frelich!and!Lorimer!1985,!Strole!and!Anderson! 1992,!Rooney!and!Waller!2003,!Wakeland!and!Swihart!2009))!hereafter!ACSA,!and!one!of! low!deer!preference!(Prunus)serotina!Ehrh.,!(Horsley!et!al.!2003,!Wakeland!and!Swihart! 2009)),!hereafter!PRSE.!!CAOV)bare!root!seedlings!were!obtained!from!Musser!Forests,! Inc.,(Indiana,!PA).)!PRSE,!ACSA!and!QUAL!bare!root!seedlings!were!obtained!from!Cold! Stream!Farm!(Free!Soil,!MI).!!All!tree!seedlings!were!12.5!–!30.5!cm!tall!when! planted.!!!!Tree!seedlings!were!randomly!assigned!to!a!cardinal!and!an!intercardinal! direction,!planting!four!trees!0.5!m!under!the!canopy!of!L.)maackii!(cardinal)!and!four!trees! 0.5!m!outside!of!the!canopy!of!L.)maackii!(intercardinal,!Figure!3).!!!Seedlings!were!planted! May!31!–!June!6,!2014,!and!treated!with!Soil!Moist!water!storing!polymer!root!gel!(Musser! Forests,!Inc.)!at!the!time!of!planting,!to!help!prevent!transplant!shock.! Seedlings!were!measured!monthly!during!the!growing!season,!and!once!every!2>3! months!during!the!winter,!from!June!2014!to!June!2015!(Appendix!Table!A1).!!During!the! growing!season,!I!recorded!survival,!height,!deer!browse,!and!number!of!leaves,!while! during!the!winter,!only!height!and!deer!browse!were!recorded.!Surviving!seedlings!were! excavated!at!the!end!of!June!2015,!separated!into!roots!and!shoots,!washed,!dried!to! constant!mass!at!70!degrees!Celsius,!and!subsequently!weighed.!!!
Data!Analysis) PRSE)seedlings!were!excluded!from!all!analyses,!due!to!high!initial!mortality!(12/40! seedlings!were!never!alive!over!the!course!of!the!study,!30%!mortality)!and!complete! mortality!by!June!2015.!!In!addition,!seedlings!that!were!dead!at!the!first!census!(leafless! two!weeks!after!planting,!and!never!leafing!out)!were!excluded!from!all!analyses,!as!I! attributed!this!to!transplant!shock.!! !For!all!other!species,!tree!seedling!survival!was!assessed!via!survival!analysis,!an! analytical!tool!that!deals!with!longevity!and!is!commonly!used!in!analyses!of!time!until!
! 49! ! death!(Fan!et!al.!!2008).!!!All!seedlings!that!were!lost!over!the!course!of!the!study!were! treated!as!dead.!!All!seedlings!surviving!at!the!termination!of!the!study!were!right> censored.!!!For!each!species,!χ2)values!were!calculated!from!observed!deaths!and)Kaplan> Meier!estimates!of!predicted!survival,!and!differences!between!treatments!were!calculated! using!the!Peto!and!Peto!modification!of!the!Gehan>Wilcoxon!test!(Pyke!and!Thompson! 1986,!Rich!et!al.!2010).!!Note!that!while!graphs!show!percent!survival,!calculations!on! survival!data!were!completed!on!mortality,!the!proportion!that!died!at!each!census.!!! Browse!status!was!analyzed!using!a!split>plot,!generalized!linear!mixed>effect! model,!modified!for!binary!data!with!a!binomial!distribution.!!The!model!was!specified! using!the!location!under!or!away!from!the!canopy!of!L.)maackii!as!a!fixed!effect!and!plot!as! the!random!effect.!!Analyses!of!browse!were!completed!on!only!seedlings!within!the!deer! access!plots,!as!the!exclosures!were!effective!in!preventing!all!browse!on!planted!seedlings.!! Significance!was!determined!by!maximum!likelihood!(Laplace!Approximation).!! Final!leaf!count!was!analyzed!using!a!split>plot,!generalized!linear!mixed>effect! model,!modified!for!count!data!with!a!Poisson!distribution.!!The!model!was!specified!using! effects!of!deer!exclosure!and!location!under!or!away!from!the!canopy!of!L.)maackii!as!fixed! effects!and!plot!as!the!random!effect.!!Significance!was!determined!by!maximum!likelihood! (Laplace!Approximation).! Change!in!height,!root!mass,!shoot!mass!and!root:shoot!ratio!were!analyzed!using! linear,!mixed>effects!model!using!the!effects!of!deer!exclosure!and!under!or!away!from!L.) maackii!canopy!as!fixed!effects!and!plot!as!a!random!effect.!!!These!analyses!excluded! seedlings!that!died!during!the!one!year!experiment.!! ! All!Analyses!were!completed!in!R!version!3.1.2!(R!Core!Team!2014)!using!the!nlme!(! change!in!height,!root!mass,!shoot!mass!and!root:shoot!ratio,!Pinheiro!et!al.!2014),!lme4! (browse!status!and!final!leaf!count,!Bates!et!al.!2014),!!and!survival!(survival!analyses,! Therneau!and!Grambsch!2000,!Therneau!2015)!packages.!!!For!data!visualization!I!used!the! ggplot2!(Wickham!2009),!and!sciplot!(Morales!et!al.!2012)!packages.!!
! 50! !
Results'
Survival!! ! ACSA!seedling!survival!was!higher!in!deer!exclosures!than!in!areas!of!deer!access!(P! =!!0.041,!Table!1,!Figure!4,!Appendix!Table!A2).!!While!there!was!no!effect!of!position!under! or!away!from!the!canopy!of!L.)maackii!on!survival!of!ACSA,!there!was!an!interaction! between!the!deer!and!L.)maackii)treatments!(P!=!0.005).!!!In!areas!of!deer!access,!there!was! much!higher!survival!under!rather!than!away!from!the!canopy!of!L.)maackii.!!In!areas!of! deer!exclosure,!survival!did!not!differ!between!honeysuckle!treatments!(Figures!5!and!6).!! Survival!of!CAOV!and!QUAL!seedlings!did!not!respond!to!deer!exclosures!or!position,!and! there!were!no!interactive!effects!(Figure!4,!Appendix!Table!A2,!Figures!A1,!A2).))!
Browse!status! In!areas!of!deer!access,!there!was!not!a!significant!effect!of!position!under!or!away! from!L.)maackii)on!browse!status!of!ACSA!or!CAOV!seedlings!(Appendix!Table!A3,!Appendix! Figure!A3).!!I!could!not!analyze!browse!status!for!QUAL!as!all!seedlings!in!deer!access!plots! except!for!one!were!browsed,!leading!to!a!singularity.!!!!
Final!Leaf!Count! ! There!were!more!leaves!on!ACSA!seedlings!under!L.)maackii)compared!to!seedlings! away!from!L.)maackii!(z!=!2.044,!P!=0.0409,!Figure!7))and!more!leaves!on!ACSA!seedlings!in! areas!of!deer!exclosure!than!deer!access!(z!=!2.156,!P!=!0.0311,!Table!2)!.!)Additionally,! there!was!an!interaction!between!deer!and!L.)maackii)seedlings!on!ACSA!seedlings!leaf! count!(z!=!>2.100,!p!=!0.036).!!ACSA!seedlings!planted!away!from!L.)maackii!had!fewer! leaves!in!areas!of!deer!access!compared!to!areas!of!deer!exclosure.!!Where!seedlings!were! planted!under!honeysuckle,!there!was!no!difference!in!leaf!count!between!areas!of!deer! access!and!deer!exclosure!(Figure!8).!!In!addition,!there!was!a!significant!effect!of!site!on! ACSA!leaf!count!(z!=!3.769,!P!–!0.0002),!with!seedlings!in!Western!Woods!averaging!more! leaves!than!seedlings!in!Kramer!Preserve.!!Final!leaf!count!of!CAOV!and!QUAL!seedlings!did! not!respond!to!deer!exclosures!or!position!under!or!away!from!L.)maackii,!and!there!were! no!interactive!effects!(Table!2,!Figure!7).!
! 51! !
Change!in!Height! ! QUAL!seedlings!had!greater!loss!of!height!in!areas!of!deer!access!than!areas!of!deer! exclosure!between!July!2014!and!July!2015!(F1,!17=!6.3315,!!P!=!0.022,!Table!3,!Figure!9);! seedlings!in!deer!exclosures!lost!less!height!than!those!in!access!plots!(>0.2!±!1.7!cm!vs.!>6.8! ±!2.0!cm!(mean!±!SE),!respectively).!!Deer!exclosure!had!no!effect!on!the!change!in!height!of! surviving!ACSA!or!CAOV!seedlings.!!!Additionally,!there!was!no!significant!effect!of!position! under!or!away!from!L.)maackii,!or!of!the!interaction!between!deer!and!L.)maackii,)on!the! change!in!height!of!QUAL,!ACSA!or!CAOV!seedlings.!!
Root!Mass,!Shoot!Mass!and!Root:Shoot!Ratio! ! Root!Mass!and!Shoot!Mass!of!ACSA,!CAOV!and!QUAL!seedlings!did!not!respond!to! deer!exclosures!or!position,!and!there!were!no!significant!interactive!effects.!There!was!an! effect!of!site!on!ACSA!root!mass!(F1,!16=!5.7486,!!P!=!0.029)!and!shoot!mass!(F1,!16=!4.8638,!! P!=!0.042),!with!greater!root!and!shoot!mass!in!Western!Woods!than!in!Kramer!Preserve.!!!
Root:shoot!ratio!of!QUAL!seedlings!was!affected!by!deer!exclosure!(F1,!17=!4.5570,!!P! =!0.048),!with!greater!root:shoot!ratio!in!areas!of!deer!access!(4.19!±!0.55,!mean!±!SE)!than! areas!of!deer!exclosure!(2.84!±!0.21).!!!!There!was!also!a!trend!for!greater!root:shoot!ratio! away!from!L.)maackii)(4.02!±!0.55)!compared!to!under!L.)maackii)(2.96!±!0.19).!However,! there!was!no!interaction!between!deer!and!L.)maackii)treatments!on!QUAL!root:shoot!ratio.!! Root:shoot!ratio!of!ACSA!and!CAOV!seedlings!did!not!respond!to!deer!exclosures!or! position!under!or!away!from!L.)maackii,!and!there!were!no!interactive!effects.!
Discussion'
Facilitation! Evidence!for!facilitation!of!tree!seedlings!by!L.)maackii)would!be!an!interaction! between!the!effects!of!position!under!or!away!from!the!canopy!of!L.)maackii)and!exposure! to!deer!access!on!tree!seedling!responses,!where!in!the!presence!of!deer!seedlings! performed!better!under!L.)maackii)(Figure!1A).!!Both!survival!and!leaf!count!of!ACSA! seedlings!showed!this!pattern.!! !! ACSA!survival!was!significantly!affected!by!the!deer!and!L.)maackii!interaction;! seedlings!planted!away!from!L.)maackii!had!much!lower!survival!in!areas!of!deer!access!
! 52! ! compared!to!areas!of!deer!exclosure.!!For!seedlings!were!planted!under!L.)maackii,!there! was!no!difference!in!survival!between!areas!of!deer!access!and!deer!exclosure.!!This! suggests!that!location!under!the!canopy!of!L.)maackii)mitigates!the!large,!negative!effect!of! exposure!to!deer!browse,!thus!facilitating!the!survival!of!ACSA.!! Final!leaf!count!of!the!surviving!ACSA!seedlings!also!supports!facilitation!by!L.) maackii.!!ACSA!seedlings!planted!away!from!L.)maackii)shrubs!had!many!fewer!leaves!in! areas!of!deer!access!when!compared!to!areas!of!deer!exclosure,!but!ACSA!seedlings!planted! under!L.)maackii)shrubs,!showed!no!difference!in!leaf!count.!!!
Competition! ! Competition!would!be!indicated!by!a!significant!L.)maackii!effect!on!tree!seedling! responses!(Figure!1B).!!I!found!limited!support!that!the!interactions!between!L.)maackii! and!tree!seedlings!are!competitive!in!nature.!ACSA!seedlings!planted!under!L.)maackii) shrubs!had!a!greater!number!of!leaves!than!those!planted!away.!!This!may!be!due!to!ACSA! increasing!leaf!production!in!response!to!shade.!Additionally,!QUAL!seedlings!under!the! canopy!of!L.)maackii)showed!lower!root:shoot!ratio!than!those!away!from!L.)maackii.))This! greater!relative!allocation!to!shoot!growth!is!a!typical!response!to!light!limitation,!and!is! evidence!for!a!competitive!shading!effect!of!L.)maackii.)Hartman!and!McCarthy!(2004)! found!significantly!greater!survival!of!all!tree!seedling!species!(Cercis)canadensis,)Cornus) florida,)Fraxinus)pennsylvanica,)Juglans)nigra,)Prunus)serotina,!and!Quercus)muhlenbergii)! with!removal!of!L.)maackii,!regardless!of!deer.!Similarly,)Gorchov!and!Trisel!(2003)!found! removal!of!L.)maackii)shoots!resulted!in!greater!survival!of!Acer)saccharum)and!Fraxinus) americana!seedlings!and!a!trend!for!increased!survival!of!Quercus)rubra!,!as!well!as!a! significantly!higher!root:shoot!ratio!of!!A.)saccharum,!but!not!on!F.)americana)or!Q.)rubra! seedlings.!
Additional!Findings! ! While!I!was!only!able!to!test!the!effects!of!location!under!or!away!from!L.)maackii!on! browse!status!of!seedlings!in!deer!access!plots,!90!±!7%!(mean!±!SE,!averaged!across!status! under!or!away!from!L.)maackii)!of!QUAL!seedlings!were!browsed!in!areas!of!deer!access.! Despite!this!high!browse,!≥!80%!survival!of!QUAL!seedlings!was!recorded!in!all!treatment! combinations.!!Others!have!also!found!this!pattern!of!very!high!survival!of!Quercus)species!
! 53! ! in!response!to!browse!(Jensen!et!al.!2012,!Kern!et!al.!2012).!!Our!study!may!have!been! limited!in!its!ability!to!detect!changes!in!QUAL!survival!by!its!one>year!duration,!as!Jensen! et!al.!(2012)!found!that!effects!of!deer!browse!on!survival!of!Quercus!species!was!not! manifest!until!the!third!year!of!browse!exposure.! Presumably!because!of!this!prevalent!browse,!mean!height!growth!of!QUAL! seedlings!was!negative!–!they!typically!were!shorter!at!the!end!of!the!year!than!when! planted.!!This!loss!of!height!was!smaller!in!areas!of!deer!exclosure!than!in!areas!of!deer! access.!!However,!there!was!no!effect!of!location!under!or!away!from!L.)maackii)on!change! in!height!of!QUAL!seedlings.!!This!is!likely!due!to!how!highly!preferred!QUAL!are!as!browse! for!deer!(Strole!and!Anderson!1992,!Wakeland!and!Swihart!2009);!deer!may!be!willing!to! put!in!more!effort!to!maneuver!under!L.)maackii)shrubs!to!access!these!highly!preferred! seedlings.!!!While!these!differences!in!change!in!height!of!QUAL!seedlings!are!likely!due!to! differences!in!browse,!they!may!also!be!due,!in!part,!to!differences!in!patterns!of!allocation! to!root!or!shoot!tissues.!!QUAL!seedlings!in!areas!of!deer!access!had!lesser!shoot!mass!(1.19! ±!0.14!g!vs.!1.44!±!0.14!g)!than!areas!of!deer!exclosure,!and!the!tallest!part!of!the!seedling! was!almost!always!browsed,!suggesting!higher!tissue!loss!to!deer!browse.!!However,!QUAL! seedlings!in!areas!of!deer!access!also!had!greater!root!mass!(4.39!±!0.49!g,!mean!±!se,!vs.! 4.06!±!0.48!g)!than!seedlings!in!areas!of!deer!exclosure,!suggesting!that!there!is!a!difference! in!allocation!patterns!between!deer!treatments.! Though!Gorchov!and!Trisel!(2003)!and!Cipollini!et!al.!(2009)!had!suggested!L.) maackii)protects!tree!seedlings!and!herbs!from!deer!browse,!this!study!did!not!find! evidence!for!an!interactive!effect!of!L.)maackii!and!deer!on!browse!incidence.!!This!may! have!been!due!a!high!number!of!seedlings!across!all!species!and!treatment!types! accumulating!at!least!one!browse!event!by!the!end!of!the!study.!!Cumulative!number!of! browse!events!per!seedling!could!be!a!more!informative!variable!that!might!exhibit!this! interactive!effect!of!L.)maackii)and!deer.!!The!protective!effects!of!other!invasive!plants!on! species!other!than!tree!seedlings!has!also!been!posited;!Waller!and!Maas!(2013)!found!that! Alliaria)petiolata)cover!enhances!growth!and!survival!of!a!highly!preferred!herb,!Uvularia) gradiflorum,!in!the!presence!of!deer.!!! In!conclusion,!I!found!more!support!for!categorizing!the!relationship!between!native! tree!seedlings!and!L.)maackii!as!facilitative,!rather!than!competitive,!in!an!area!of!high!deer!
! 54! ! density.!This!facilitation!by!L.)maackii)on!ACSA!growth!and!survival!was!observed!after! only!one!year,!suggesting!that!ACSA!may!benefit!greatly!over!several!years.!!!This!benefit! may!extend!to!other!moderately!preferred!tree!species,!and!would!be!expected!to!result!in! greater!regeneration!and!recruitment!into!the!canopy.!!Differential!survivorship!of!canopy! tree!seedlings,!even!at!low!deer!densities,!may!affect!the!future!composition!of!the!forest! canopy!(Aronson!and!Handel!2011).!The!negative!effects!of!L.)maackii)on!tree!seedling! growth!are!ameliorated!by!its!mitigation!of!the!negative!effects!of!deer!browse.!In!forests! with!dense!deer!populations,!removal!of!invasive!shrubs,!such!as!L.)maackii,!that!have!this! protective!effect!would!be!expected!to!reduce!regeneration!of!some!tree!species.!! ! !
! 55! !
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Horsley,!S.!B.,!S.!L.!Stout,!and!D.!S.!deCalesta.!2003.!White>tailed!deer!impact!on!the! vegetation!dynamics!of!a!northern!hardwood!forest.!Ecological!Applications!13:98– 118.! Jensen,!A.!M.,!F.!Götmark,!and!M.!Löf.!2012.!Shrubs!protect!oak!seedlings!against!ungulate! browsing!in!temperate!broadleaved!forests!of!conservation!interest:!a!field! experiment.!Forest!Ecology!and!Management!266:187–193.! Kern,!C.!C.,!P.!B.!Reich,!R.!A.!Montgomery,!and!T.!F.!Strong.!2012.!Do!deer!and!shrubs! override!canopy!gap!size!effects!on!growth!and!survival!of!yellow!birch,!northern! red!oak,!eastern!white!pine,!and!eastern!hemlock!seedlings?!Forest!Ecology!and! Management!267:134–143.! Lieurance,!D.,!and!D.!Cipollini.!2012.!Damage!levels!from!arthropod!herbivores!on!Lonicera) maackii!suggest!enemy!release!in!its!introduced!range.!Biological!Invasions!14:863– 873.! Lieurance,!D.,!and!D.!Cipollini.!2013.!Exotic!Lonicera!species!both!escape!and!resist! specialist!and!generalist!herbivores!in!the!introduced!range!in!North!America.! Biological!invasions!15:1713–1724.! Luken,!J.!O.,!and!J.!W.!Thieret.!1996.!Amur!honeysuckle,!its!fall!from!grace.!BioScience! 46:18–24.! McShea,!W.!J.!2012.!Ecology!and!management!of!white>tailed!deer!in!a!changing!world.! Annals!of!the!New!York!Academy!of!Sciences!1249:45–56.! Merriam,!R.!W.,!and!E.!Feil.!2002.!The!potential!impact!of!an!introduced!shrub!on!native! plant!diversity!and!forest!regeneration.!Biological!Invasions!4:369–373.! Miller,!J.!H.!2006.!Nonnative!invasive!plants!of!southern!forests.!USDA!Forest!Service/UNL! Faculty!Publications:103.! Morales,!M.,!!with!code!developed!by!the!R.!D.!C.!Team,!!with!general!advice!from!the!R.!> help.!listserv!community,!and!!especially!D.!Murdoch.!2012.!sciplot:!scientific! graphing!functions!for!factorial!designs.! Perea,!R.,!and!L.!Gil.!2014.!Tree!regeneration!under!high!levels!of!wild!ungulates:!The!use!of! chemically!vs.!physically>defended!shrubs.!Forest!Ecology!and!Management!312:47– 54.!
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Pergams,!O.!R.!W.,!and!J.!E.!Norton.!2006.!Treating!a!single!stem!can!kill!the!whole!shrub:!a! scientific!assessment!of!buckthorn!control!methods.!Natural!Areas!Journal!26:300– 309.! Pinheiro,!J.,!D.!Bates,!S.!DebRoy,!D.!Sarkar,!and!R!Core!Team.!2014.!nlme:!linear!and! nonlinear!mixed!effects!models.! Pyke,!D.!A.,!and!J.!N.!Thompson.!1986.!Statistical!analysis!of!survival!and!removal!rate! experiments.!Ecology:240–245.! R!Core!Team.!2014.!R:!a!language!and!environment!for!statistical!computing.!R!Foundation! for!Statistical!Computing,!Vienna,!Austria.! Rich,!J.!T.,!J.!G.!Neely,!R.!C.!Paniello,!C.!C.!J.!Voelker,!B.!Nussenbaum,!and!E.!W.!Wang.!2010.!A! practical!guide!to!understanding!Kaplan>Meier!curves.!Otolaryngology!143:331– 336.! Rooney,!T.!2009.!High!white>tailed!deer!densities!benefit!graminoids!and!contribute!to! biotic!homogenization!of!forest!ground>layer!vegetation.!Plant!Ecology!202:103– 111.! Rooney,!T.!P.,!and!D.!M.!Waller.!2003.!Direct!and!indirect!effects!of!white>tailed!deer!in! forest!ecosystems.!Forest!Ecology!and!Management!181:165–176.! Smit,!C.,!C.!Vandenberghe,!J.!Ouden,!and!H.!Müller>Schärer.!2007.!Nurse!plants,!tree!saplings! and!grazing!pressure:!changes!in!facilitation!along!a!biotic!environmental!gradient.! Oecologia!152:265–273.! Strole,!T.!A.,!and!R.!C.!Anderson.!1992.!White>tailed!deer!browsing:!species!preferences!and! implications!for!central!Illinois!forests.!Natural!Areas!Journal!12:139–144.! Therneau,!T.!M.!2015.!A!package!for!survival!analysis!in!S.! Therneau,!T.!M.,!and!P.!M.!Grambsch.!2000.!Modeling!survival!data:!extending!the!cox! model.!Springer,!New!York.! Vandenberghe,!C.,!C.!Smit,!M.!Pohl,!A.!Buttler,!and!F.!Freléchoux.!2009.!Does!the!strength!of! facilitation!by!nurse!shrubs!depend!on!grazing!resistance!of!tree!saplings?!Basic!and! Applied!Ecology!10:427–436.! Wakeland,!B.,!and!R.!K.!Swihart.!2009.!Ratings!of!white>tailed!deer!preferences!for!woody! browse!in!Indiana.!Pages!96–101!Proceedings!of!the!Indiana!Academy!of!Science.!
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Waller,!D.!M.,!and!L.!I.!Maas.!2013.!Do!white>tailed!deer!and!the!exotic!plant!garlic!mustard! interact!to!affect!the!growth!and!persistence!of!native!forest!plants?!Forest!Ecology! and!Management!304:296–302.! Webster,!C.!R.,!M.!A.!Jenkins,!and!S.!Jose.!2006.!Woody!invaders!and!the!challenges!they! pose!to!forest!ecosystems!in!the!eastern!United!States.!Journal!of!Forestry!104:366– 374.! Wickham,!H.!2009.!ggplot2:!elegant!graphics!for!data!analysis.!Springer!New!York.! Wiegmann,!S.!M.,!and!D.!M.!Waller.!2006.!Fifty!years!of!change!in!northern!upland!forest! understories:!Identity!and!traits!of!“winner”!and!“loser”!plant!species.!Biological! Conservation!129:109–123.! Woods,!K.!D.!1993.!Effects!of!invasion!by!Lonicera)tatarica!L.!on!herbs!and!tree!seedlings!in! four!New!England!forests.!American!Midland!Naturalist:62–74.! !
! !
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Table'1.!!ACSA!seedling!survival!by!treatment!type.!X2)values!were!calculated!from! observed!survival!and)Kaplan>Meier!estimates!of!predicted!survival,!and!differences! between!treatments!were!calculated!using!the!Peto!and!Peto!modification!of!the!Gehan> Wilcoxon!test.!!Significant!effects!are!bolded.! ! Treatment! Chisq! df! P! Deer! 4.2! 1! 0.0407! Honeysuckle! 0.7! 1! 0.409! Deer:Honeysuckle! 12.7! 3! 0.00545! ! ! !
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Table'2.!Significance!of!treatment!effects!on!leaf!count!of!surviving!seedlings!of!each! species.!Significance!based!on!a!split>plot,!generalized!linear!mixed>effect!model,!modified! for!count!data!with!a!Poisson!distribution.!Significance!of!effects!were!determined!by! maximum!likelihood!(Laplace!Approximation).!!Significant!effects!are!bolded.! ! Response! Species!! Treatment! Estimate! Std.!Error! z!value! Pr(>|z|)! Leaf! Count! ACSA! Site! ' 1.035! ' 0.275! ' 3.7690! ' 0.0002! ' ! ! Deer' 1.620' 0.752' 2.1560' 0.0311' ! ! Honeysuckle' 1.527' 0.747' 2.0440' 0.0409' ! ! Deer*Honeysuckle' >1.671' 0.796' >2.1000' 0.0357' ! CAOV! ! ! Site! ! >0.243! ! 0.302! ! >0.8040! ! 0.4214! ! ! ! Deer! >0.229! 0.300! >0.7640! 0.4446! ! ! Honeysuckle! >0.117! 0.281! >0.4170! 0.6766! ! CAOV! ! ! Site! ! 0.216! ! 0.161! ! 1.3420! ! 0.18! 00! ! ! Deer! 0.173! 0.160! 1.0810! 0.2790! !! !!! Honeysuckle! 0.061! 0.111! 0.5490! 0.5830! ! ! ! ! ! ! ! ! ! !
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Table'3.!!Significance!of!treatment!effects!on!change!in!height!of!surviving!seedlings!of! each!species.!!Values!determined!using!split>plot!analysis!of!variance.!!Significant!effects! are!bolded.!! ! Response! Species! Treatment! df) den!df! F! P! Change!in! Height! ACSA! Site! ! 1)! 16! ! 1.4470! ! 0.2465! ! ! ! Deer! 1! 16! 0.3228! 0.5778! ! ! Honeysuckle! 1! 8! 0.1190! 0.7391! ! CAOV! ! ! Site! ! 1!! 15! ! 3.2739! ! 0.0905! ! ! ! Deer! 1! 15! 0.0182! 0.8945! ! ! Honeysuckle! 1! 7! 0.1388! 0.7205! ! QUAL! ! ! Site! ! 1!! 17! ! 0.6607! ! 0.4276! ! ! ! Deer' 1' 17' 6.3314' 0.0222' ' ' Honeysuckle! 1! 17! 1.0680! 0.3159! ! ! ! !
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Facilitation Competition 100 100 A) B) Deer ACC EXC 75 75
50 50 Response Response
25 25
0 0
Away Under Away Under Honeysuckle Honeysuckle ! ! Figure'1.!!Predictions!of!tree!seedling!responses!(survival,!growth,!etc.).!!Predictions!based! on!the!hypotheses!of!A)!facilitative!interactions,!or!B)!competitive!interactions,!between! Lonicera)maackii!and!native!tree!seedlings!under!and!away!from!the!canopy!of!L.)maackii! and!with!(EXC)!and!without!(ACC)!deer!exclosures!between!L.)maackii.!!! ! !
! 64! !
! Figure'2.!Schematic!of!study!design.!!Tree!seedlings!were!planted!at!each!of!the!cardinal! directions!under!the!canopy!of!Lonicera)maackii!and!at!each!of!the!intercardinal!directions! away!from!the!canopy!of!L.)maackii.)Each!L.)maackii!shrub!was!randomly!assigned!to!have!a! deer!exclosure!or!not.! ! !
! 65! !
! ' Figure'3.!!Map!of!study!locations!within!the!Miami!University!Natural!Areas.!!Pink!dots! denote!areas!of!deer!access!plots,!while!yellow!dots!denote!areas!of!deer!exclosure,!for!a! total!of!20!plots!across!2!natural!areas.!!The!orange!line!marks!the!boundary!of!the!natural! areas.!!Natural!Areas!are!annotated!with!their!names.
! 66! !
ACSA CAOV QUAL
100
75
Deer
50 Access Exclosure Survival (Percent) Survival
25
0
Away Under Away Under Away Under Honeysuckle Figure'4.!!!Percent!survival!(+/3!SE)!of!seedlings.!!Percent!survival!of!Acer%saccharum%(ACSA),!Carya%ovata!(CAOV),!and!Quercus% alba!(QUAL)!seedlings!planted!under!or!away!from!the!canopy!of!L.%maackii!and!with!(ACC)!or!without!(EXC)!deer!access.!!!
67! !
Figure'5.!Survival!curves!for!Acer%saccharum!(ACSA).!!Survival!curves!for!ACSA%in!each!treatment!over!the!course!of!the!50! week!experiment.
68! !
! Figure'6.!!Interaction!plot!of!Acer%saccharum!survival,!mean!±!SE!for!each!treatment.!!Solid! circles!and!lines!represent!areas!of!deer!access,!open!circles!and!dotted!lines!represent! areas!of!deer!exclosure.!!Location!under!or!away!from!the!canopy!of!L.%maackii!is!shown!on! the!x?axis.!! ! !
69! !
! Figure'7.!!!Final!leaf!count!of!seedlings.!!Boxplots!of!final!leaf!count!of!Acer%saccharum% (ACSA),!Carya%ovata!(CAOV),!and!Quercus%alba!(QUAL)!seedlings!planted!under!or!away! from!the!canopy!of!L.%maackii!and!with!(ACC)!or!without!(EXC)!deer!access.!!! ! ! !
70! !
! Figure'8.!!Interaction!plot!of!Acer%saccharum!seedling!leaf!count,!mean!±!SE!for!each! treatment.!!Solid!circles!and!lines!represent!areas!of!deer!access,!open!circles!and!dotted! lines!represent!areas!of!deer!exclosure.!!Location!under!or!away!from!the!canopy!of!L.% maackii!is!shown!on!the!x?axis.!! ! !
71! !
' Figure'9.!Change!in!height!between!July!2014!and!June!2015!of!seedlings.!!Boxplots!of! change!in!height!for!Acer%saccharum%(ACSA),!Carya%ovata!(CAOV),!and!Quercus%alba%(QUAL)! seedlings!planted!under!or!away!from!the!canopy!of!L.%maackii!and!with!(ACC)!or!without! (EXC)!deer!access.!!Values!were!determined!from!the!final!height!(June!2015)!minus!the! initial!height!(July!2014).!!! ! !
72! !
Chapter'4.'Browse'by'White:tailed'Deer'Decreases'Cover'and'Growth'of' the'Invasive'Shrub,'Lonicera)maackii)) ! Note:!Two!data!sets!incorporated!into!this!chapter!were!obtained!by!undergraduate! collaborator!Christina!M.!Haffey;!the!2015!Lonicera%maackii%basal!areas!and!browse! damage,!Ms.!Haffey!will!be!a!co?author!when!this!chapter!is!submitted!for!publication.!! !
Introduction'' The!success!of!non?native!species!in!their!introduced!range!is!often!related!to! release!from!consumptive!pressure!from!native!enemies!(Enemy!Release!Hypothesis,!ERH,! Keane!and!Crawley!2002)!While!this!hypothesis!primarily!addresses!the!success!of! introduced!plant!species,!it!also!makes!predictions!about!the!interactions!between! generalist!herbivores!and!introduced!plant!species.!!ERH!predicts!that!generalist! herbivores!will!avoid!browsing!invasive!species!in!preference!of!natives,!promoting!the! success!of!invasive!species!in!their!introduced!range!(Keane!and!Crawley!2002,!Colautti!et! al.2004).!! Conversely,!the!biotic!resistance!hypothesis,!BRH,!first!articulated!by!Elton!(1958),! suggests!that!native!species,!such!as!generalist!herbivores,!can!function!to!suppress!an! invasive!plant’s!establishment!through!the!exploitation!of!novel!food!sources!(Verhoeven! et!al.!2009,!Morrison!and!Hay!2011).!!This!suppressive!effect!has!been!shown!between! many!pairs!of!native!enemies!and!invasive!plants!(Parker!et!al.!2006a,!2006b,!Ricciardi!and! Ward!2006),!with!native!herbivores!suppressing!invasive!plant!species!far!more!often!than! native!plant!species!(Parker!et!al.!2006a).!However,!Parker!et!al.'s!(2006a)!review!included! no!studies!from!temperate!deciduous!forest,!so!the!effects!of!native!herbivores!on!invasive! plants!in!these!forests!are!as!of!yet!poorly!understood.! We!investigated!whether!herbivory!by!a!generalist!herbivore!on!a!non?native!shrub! invasive!in!deciduous!forest!is!consistent!with!predictions!of!ERH!or!BRH.%Lonicera%maackii% (Rupr.)!Herder!(Caprifoliaceae),!Amur!honeysuckle,!is!a!large!shrub!that!is!native!to! northeast!Asia!(Luken!and!Thieret!1996),!and!one!of!several!bush!honeysuckles!invasive!in! the!eastern!United!States.!The!current!distribution!of!L.%maackii%spans!the!eastern!half!of!
73! ! the!United!States,!and!this!species!is!considered!invasive!and!regulated!in!eight!states! (EDDMapS!2015).!The!success!of!L.%maackii%in!North!America!has!been!attributed!to!many! factors,!including!phenotypic!plasticity!(Luken!et!al.!1995,!1997)!early!leaf!expansion! (McEwan!et!al.!2009),!late!leaf!retention!(Wilfong!et!al.!2009),!high!freeze!tolerance!of! leaves!(McEwan!et!al.!2009),!and!the!absence!of!specialist!herbivores!(Lieurance!and! Cipollini!2011).!!Lonicera%maackii%decreases!growth!and!survival!of!tree!seedlings!(Gorchov! and!Trisel!2003),!survival!and!reproduction!of!forest!annuals!(Gould!and!Gorchov!2000),! and!growth!and!reproduction!of!perennial!herbs!(Miller!and!Gorchov!2004).!!The! successful!invasion!of!many!eastern!North!American!forests!by!Lonicera%maackii%has!been! attributed,!at!least!in!part,!to!ERH,!based!on!very!low!levels!of!arthropod!herbivory! (Lieurance!and!Cipollini!2011).!!Herbivory!by!vertebrate!herbivores,!however,!has!not!yet! been!reported!for!this!shrub!in!its!introduced!range.!!! A!native!generalist!herbivore!that!is!potentially!very!important!to!the!invasion!of! this!and!other!non?native!plants!is!White?tailed!deer,!Odocoileus%virginianus!Zimmerman,! hereafter!deer.!Deer!are!notorious!generalists,!consuming!several!hundred!different! species!of!algae,!fungi,!herbs,!shrubs,!and!trees!(Atwood!1941).!!Deer!densities!in!many! parts!of!the!United!States!are!well!above!historical!levels,!often!30?50!deer/km2!where!not! managed!(McShea!2012),!causing!negative!impacts!on!tree!regeneration!(Horsley!et!al.! 2003,!Côté!et!al.!2004).!!At!even!moderate!densities,!deer!greatly!alter!structure!and! composition!of!forests!(Rooney!2009).!!At!densities!as!low!as!3?10!deer/km2!deer! negatively!impact!preferred!browse!species!(McShea!2012).!!In!some!systems,!deer! increase!the!prevalence!of!invasive!plant!species!by!selectively!feeding!on!native!plant! species!(Knight!et!al.!2009).!!While!the!browse!preference!by!deer!of!L.%maackii%is!not!well! studied,!its!leaves!have!low!palatability!for!insects!(Lieurance!and!Cipollini!2012,!2013),!! and!the!congeneric!invasive!shrub!L.%morrowii%A.!Gray,%is!of!moderate!preference!to!deer! (Averill!et!al.!2016).! We!investigated!whether!deer!herbivory!on!L.%maackii%is!high!or!low,!and!thus! consistent!with!the!ERH!or!the!BRH.!
74! !
Methods' ''
Field!Methods! !!!!!!!!!!!This!study!utilized!deer!exclosures!and!paired!deer!access!plots!in!five!sites!in!the! Miami!University!Natural!Areas,!Butler!County,!Ohio!(39°!29’!?!39°!31’!N,!84°!42’!?!84°!43’! W).!!Sites!were!separated!by!≥!1!km!and!chosen!to!have!(1)!a!similar,!moderate!level!of! invasion!by!L.%maackii%(stem!basal!area!(BA)!0.58!–!1.57!m2/ha),!!(2)!closed?canopy! deciduous!forest,!and!(3)!level!topography.!Deer!densities!in!this!area!range!from!11.18!±! 0.644!(Mean!±!!SE)!deer!per!km2!in!the!summer!to!15.42!±!4.406!deer!per!km2!in!the!winter! (Barrett!2014).!!Within!each!site,!two!20m?by?20m!plots!that!were!very!similar!were! selected!and!randomly!assigned!to!deer!exclosure!and!deer!access!treatments.!Three!meter! tall!fencing!was!placed!around!the!deer!exclosure!plots!in!summer!2010.!Half!plots!were! randomly!assigned!to!be!L.%maackii%removed!or!intact.!!Only!the!half!of!each!deer!exclosure! and!deer!access!plot!with!L.%maackii%left!intact!were!assessed!in!this!study;!these!are! hereafter!referred!to!as!‘plots.’!!! The!BA!of!L.%maackii%in!each!plot!was!determined!in!Oct.!2010,!shortly!after! exclosures!were!constructed,!and!again!in!May?June!2015.!!At!each!census!we!measured!the! basal!diameter!of!every!live!L.%maackii%stem!≥!0.1!cm!and!used!these!diameters!to!calculate! each!plot’s!BA.! To!estimate!the!leaf!frequency!of!L.%maackii%we!sampled!18!points!(every!2!m!along! two!20!m!transects)!in!each!plot.!!At!each!sample!point,!we!sampled!a!1000cm2!(20cm?by? 50cm)!subplot!at!two!dates!during!the!2014!growing!season,!using!two!different!methods.!! Between!June!12!and!June!16,!2014,!we!scored!presence!or!absence!of!L.%maackii%leaves!in! each!subplot!at!each!of!four!heights!(0.5,!1,!1.5!and!2!meters!from!the!ground),!chosen!to! span!the!range!of!heights!at!which!deer!browse!occurs!(between!0.3!and!2!m,!Rooney!and! Waller!2003).!Between!July!16!and!July!22,!2014,!we!recorded!the!presence/absence!of!L.% maackii%leaves!in!each!subplot!within!each!of!three!height!ranges,!0!?!0.5!m,!0.5!?!1!m!and!1! ?!1.5!meters.!!! To!estimate!the!extent!of!deer!browse!on%L.%maackii%we!surveyed!for!browse!on! twigs!along!two!20!m!x!1!m!belt!transects!in!each!plot!between!May!19!and!June!10,!2015.!! For!each!L%maackii%plant!rooted!within!a!transect,!we!examined!first?year!twigs!in!the!30!–!
75! !
200!cm!height!range,!beginning!with!the!twig!closest!to!the!transect!and!proceeding!first! distal!to!the!starting!twig,!then!proximal.!Up!to!10!twigs!per!plant!were!examined!and! scored!based!on!browse!status.!!!!
Data!Analysis! We!tested!whether!initial!(2010)!L.%maackii%BA!differed!between!deer!exclosure!and! access!plots!with!a!paired!two?way!Student's!t?test.!!We!tested!whether!deer!exclosure! resulted!in!higher!L.%maackii%foliage!frequency!at!each!height!(June!2014)!and!in!each!of!the! height!ranges!(July!2014)!and!higher!L.%maackii%BA!(2015)!and!BA!growth!(BA2015!?!BA2010!)! using!paired!one?way!Student’s!t?tests,!after!checking!assumptions!for!equal!variances! using!Levene’s!tests.!!!All!analyses!were!completed!using!the!stats!(R!Core!Team!2014)!and! car!(Fox!and!Weisberg!2011)!packages!within!R,!version!3.1.2!(R!Core!Team!2014).!!
Results' ! Lonicera%maackii%basal!area!did!not!differ!between!deer!exclosure!and!access! treatments!when!exclosures!were!first!installed!(2010;!P!=!0.988),!and!BA!was!not!greater! in!exclosures!in!2015!(P!=!0.208).!!However,!there!was!greater!basal!area!growth!from! 2010!to!2015!in!exclosures!(t!=!?2.2343,!df!=!4,!P!=!0.045!Figure!1).! % The!proportion!of!first?year!L.%maackii%twigs!with!evidence!of!herbivore!browse! ranged!from!1.66!%!to!5.61!%!(3.29!±!0.91!%,!Mean!±!SE)!among!the!five!deer!access!plots! (Table!1).!!! ! There!was!higher!frequency!of!L.%maackii%leaves!at!0.5!m!(P!=!0.042!and!1!m!(P!=! 0.004)!in!deer!access!plots!compared!to!deer!exclosure!plots!(Figure!2,!Appendix!Table! A1).!!Exclosure!plots!did!not!have!greater!L.%maackii%leaf!frequency!at!1.5!m!(P!=!0.414)!or!2! m!(P!=!0.749).!%Deer!exclosure!plots!had!higher!L.%maackii%leaf!frequency!than!deer!access! plots!in!the!height!ranges!of!0.5!?!1!m!and!1!?!1.5!m!(P!=!0.024!and!P!=!0.002,!respectively,! Figure!3,!Appendix!Table!A1).!There!was!not!greater!L.%maackii%leaf!frequency!in!deer! exclosure!plots!in!the!0!?!0.5!m!height!range!(P!=!0.213).!
Discussion' ! The!findings!of!this!study!support!the!prediction!stemming!from!the!BRH,!that! native!species!suppress!an!invasive!plant!through!herbivory.!!Our!findings!contrast!with!
76! ! the!ERH,!which!predicts!little!or!no!herbivory!of!generalists!on!invasive!plants.!!However,! this!study!does!not!address!whether!deer!prefer!L.%maackii%to!native!species.!Nevertheless,! it!does!show!that!their!browse!is!strongly!impacting!cover!and!architecture!of!an!invasive! shrub.!! !We!not!only!documented!that!deer!browse!on!L.%maackii%twigs,!but!present! evidence!this!browse!is!sufficient!to!change!shrub!architecture!and!reduce!growth.!!Where! deer!were!present,!leaf!cover!of!L.%maackii%was!reduced!in!the!middle!portion!(1!–!1.5!m)!of! their!typical!browsing!height!range!(0.3!–!2!m,!Rooney!and!Waller!2003),!while!these! reductions!were!not!present!below!0.5!m!or!above!1.5!m.!!Basal!area!growth!of!L.%maackii% over!the!five?year!period!was!greater!in!plots!where!deer!were!excluded,!both!overall,!and! at!four!of!the!five!sites.!!The!fifth!site!(College!Woods),!where!growth!did!not!differ,!appears! to!be!a!site!of!relatively!low!deer!abundance!and!browse!pressure.!!This!site!had!low! browse!damage!on!new!L.%maackii%twigs!(Table!1)!and!the!lowest!winter!deer!presence! estimate!of!any!of!the!sites!(6.3!deer!per!km2,!Barrett!2014).! The!strong!effects!of!deer!on!L.%maackii!leaf!frequency!combined!with!the!modest! effects!on!BA!indicate!that!deer!impact!the!architecture!of!this!invasive!shrub!more!than!its! growth.!!The!largest!L.%maackii%shrubs!in!each!plot!were!taller!than!2!m,!and!therefore!part! of!their!crowns!were!inaccessible!to!deer!and!capable!of!supporting!growth.!!! While!we!cannot!be!certain!that!all!twigs!scored!as!browsed!were!browsed!by!deer,! and!not!rabbits!or!other!herbivores,!most!showed!the!classic!shredded!stem!characteristic! of!deer!browse.!!Additionally,!we!have!directly!observed!deer!browsing!L.%maackii%in!the! study!area!and!elsewhere,!and!Guiden!et!al.!(2015)!found!that!62!±!14%!of!fruiting!L.% maackii%branches!in!nearby!forest!edge!habitat!showed!evidence!of!deer!browse!during!a!3! month!period!from!mid?fall!to!mid?winter,!2012?2013.!!While!the!browse!rate!we!report! here!may!appear!low!(mean!of!3.29%!across!sites),!this!corresponds!to!only!browse!on! first?year!twigs!between!bud!break!in!early!spring!and!our!survey!in!late!spring,!and! cumulative!annual!browse!should!be!greater.!Herbivory!on!this!invasive!with!early!leaf! expansion!(McEwan!et!al.!2009)!may!be!particularly!great!early!in!spring!when!protein! needs!of!deer!are!high!and!other,!native!woody!plants!are!leafless%(Smith!2013).!! Alternatively,!deer!browse!pressure!on!L.%maackii%may!be!low!in!spring,!a!period!when! higher!preference!species,!such!as!many!spring!ephemerals,!are!available.!!A!survey!to!
77! ! quantify!the!annual!pattern!of!deer!browse!on!L.%maackii%in!this!area!has!been!initiated!(K.! Martinod,!unpubl.!Data).! ! This!study!is!the!first!to!show!that!browse!by!deer!is!sufficient!to!reduce!the!cover!of! L.%maackii!in!eastern!U.S.!forests,!although!the!congeneric!invasive!vine,!L.%japonica,!has! long!been!planted!as!deer!forage!(Stransky!1984).!!Deer!exclosure!was!reported!to!result!in! higher!prevalence!of!stems!of!another!invasive!bush!honeysuckle,!Lonicera%tartarica!L.,! after!8!years!of!succession!in!an!old!field!(Bowers!1997).!!The!differences!in!L.%maackii% cover!and!BA!reported!here!manifested!after!only!4!and!5!years!of!deer!exclosure,! respectively.!!That!L.%maackii%has!grown!to!be!the!dominant!shrub!in!these!stands!and! others!in!the!Midwest!may!be!a!result!of!deer!initially!failing!to!recognize!L.%maackii%as!food! source!(Morrison!and!Hay!2011).!!Deer!now!seem!to!have!overcome!that!naivety,! exploiting%L.%maackii%for!browse.!Lonicera%maackii%may!be!an!important!food!source!for! deer!in!areas!where!this!shrub!is!particularly!abundant!and/or!where!more!preferred! woody!browse!species!have!become!scarce!due!to!years!of!herbivory!by!overly!abundant! deer.!!! The!effect!of!deer!on!invasive!plant!cover!may!have!important!implications!for! understanding!the!combined!effects!of!deer!and!invasive!plants!on!native!plants.!!Our! finding!that!deer!reduce!L.%maackii%leaf!frequency!suggests!they!mitigate!the!negative! effects!of!L.%maackii%on!native!plants!(See!chapter!2).!!!Where!this!is!the!case,!the!negative! effects!of!deer!and!invasive!plants!would!not!be!additive,!i.e.!the!combined!negative!effect! on!natives!would!be!less!than!expected!from!the!individual!negative!effects.!!This!statistical! interaction!has!been!found!both!for!spring!perennial!herb!abundance!and!Maianthemum% racemosum!L.!leaf!number!in!a!recent!experiment!that!combined!deer!exclosure!and!L.% maackii!removal!(Christopher!et!al.!2014).!!Similarly,!I!found!the!negative!effect!of!L.% maackii!on!tree!seedling!cover!and!summer!herb?layer!species!richness!was!only!manifest! where!deer!were!excluded!(see!chapter!2),!suggesting!that!deer!browse!on!L.%maackii%is! sufficient!to!reduce!its!cover!and!shading!of!the!forest!floor,!therefore!reducing!the! negative!impact!of!L.%maackii!on!tree!seedling!cover!and!plant!species!richness.!At!a!larger! scale,!deer!consumption!of!invasive!shrubs!may!inflate!the!population!density!of!this! generalist!herbivore,!which!in!turn!may!suppress!native!plants,!an!example!of!apparent! competition!(Smith!2013).!! !
78! !
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Gorchov,!D.!L.,!and!D.!E.!Trisel.!2003.!Competitive!effects!of!the!invasive!shrub,!Lonicera% maackii%(Rupr.)!Herder!(Caprifoliaceae),!on!the!growth!and!survival!of!native!tree! seedlings.!Plant!Ecology!166:13–24.! Gould,!A.!M.!A.,!and!D.!L.!Gorchov.!2000.!Effects!of!the!exotic!invasive!shrub!Lonicera% maackii%on!the!survival!and!fecundity!of!three!species!of!native!annuals.!The! American!Midland!Naturalist!144:36–50.! Guiden,!P.,!D.!Gorchov,!C.!Nielsen,!and!E.!Schauber.!2015.!Seed!dispersal!of!an!invasive! shrub,!Amur!honeysuckle!(Lonicera%maackii),!by!white?tailed!deer!in!a!fragmented! agricultural?forest!matrix.!Plant!Ecology!216:939–950.! Horsley,!S.!B.,!S.!L.!Stout,!and!D.!S.!deCalesta.!2003.!White?tailed!deer!impact!on!the! vegetation!dynamics!of!a!northern!hardwood!forest.!Ecological!Applications!13:98– 118.! Keane,!R.!M.,!and!M.!J.!Crawley.!2002.!Exotic!plant!invasions!and!the!enemy!release! hypothesis.!Trends!in!Ecology!&!Evolution!17:164–170.! Knight,!T.!M.,!J.!L.!Dunn,!L.!A.!Smith,!J.!Davis,!and!S.!Kalisz.!2009.!Deer!facilitate!invasive! plant!success!in!a!Pennsylvania!forest!understory.!Natural!Areas!Journal!29:110– 116.! Lieurance,!D.,!and!D.!Cipollini.!2011.!Damage!levels!from!arthropod!herbivores!on!Lonicera% maackii%suggest!enemy!release!in!its!introduced!range.!Biological!Invasions!14:863– 873.! Luken,!J.!O.,!L.!M.!Kuddes,!and!T.!C.!Tholemeier.!1997.!Response!of!understory!species!to! gap!formation!and!soil!disturbance!in!Lonicera%maackii%thickets.!Restoration!Ecology! 5:229–235.! Luken,!J.!O.,!and!J.!W.!Thieret.!1996.!Amur!honeysuckle,!its!fall!from!grace.!BioScience! 46:18–24.! Luken,!J.!O.,!T.!C.!Tholemeier,!L.!M.!Kuddes,!and!B.!A.!Kunkel.!1995.!Performance,!plasticity,! and!acclimation!of!the!nonindigenous!shrub!Lonicera%maackii%(Caprifoliaceae)!in! contrasting!light!environments.!Canadian!Journal!of!Botany!73:1953–1961.! McEwan,!R.!W.,!M.!K.!Birchfield,!A.!Schoergendorfer,!and!M.!A.!Arthur.!2009.!Leaf!phenology! and!freeze!tolerance!of!the!invasive!shrub!Amur!honeysuckle!and!potential!native! competitors.!The!Journal!of!the!Torrey!Botanical!Society!136:212–220.!
80! !
McShea,!W.!J.!2012.!Ecology!and!management!of!white?tailed!deer!in!a!changing!world.! Annals!of!the!New!York!Academy!of!Sciences!1249:45–56.! Miller,!K.!E.,!and!D.!L.!Gorchov.!2004.!The!invasive!shrub,!Lonicera%maackii,!reduces!growth! and!fecundity!of!perennial!forest!herbs.!Oecologia!139:359–375.! Morrison,!W.!E.,!and!M.!E.!Hay.!2011.!Herbivore!preference!for!native!vs.!exotic!plants:! generalist!herbivores!from!multiple!continents!prefer!exotic!plants!that!are! evolutionarily!naïve.!PLoS!ONE!6:e17227.! Parker,!J.!D.,!D.!E.!Burkepile,!and!M.!E.!Hay.!2006a.!Opposing!effects!of!native!and!exotic! herbivores!on!plant!Invasions.!Science!311:1459–1461.! Parker,!J.!D.,!D.!E.!Burkepile,!and!M.!E.!Hay.!2006b.!Response!to!comment!on!“opposing! effects!of!native!and!exotic!herbivores!on!plant!invasions.”!Science!313:298–298.! R!Core!Team.!2014.!R:!a!language!and!environment!for!statistical!computing.!R!Foundation! for!Statistical!Computing,!Vienna,!Austria.! Ricciardi,!A.,!and!J.!M.!Ward.!2006.!Comment!on!“opposing!effects!of!native!and!exotic! herbivores!on!plant!invasions.”!Science!313:298–298.! Rooney,!T.!2009.!High!white?tailed!deer!densities!benefit!graminoids!and!contribute!to! biotic!homogenization!of!forest!ground?layer!vegetation.!Plant!Ecology!202:103– 111.! Rooney,!T.!P.,!and!D.!M.!Waller.!2003.!Direct!and!indirect!effects!of!white?tailed!deer!in! forest!ecosystems.!Forest!Ecology!and!Management!181:165–176.! Siemann,!E.,!and!W.!Rogers.!2003.!Reduced!resistance!of!invasive!varieties!of!the!alien!tree! Sapium%sebiferum!to!a!generalist!herbivore.!Oecologia!135:451–457.! Siemann,!E.,!and!W.!E.!Rogers.!2001.!Genetic!differences!in!growth!of!an!invasive!tree! species.!Ecology!Letters!4:514–518.! Smith,!L.!M.!2013.!Extended!leaf!phenology!in!deciduous!forest!invaders:!mechanisms!of! impact!on!native!communities.!Journal!of!Vegetation!Science!24:979–987.! Stransky,!J.!J.!1984.!Forage!yield!of!Japanese!honeysuckle!after!repeated!burning!or! mowing.!Journal!of!Range!Management!37:237!–!238.! Verhoeven,!K.!J.!F.,!A.!Biere,!J.!A.!Harvey,!and!W.!H.!Van!Der!Putten.!2009.!Plant!invaders!and! their!novel!natural!enemies:!who!is!naïve?!Ecology!Letters!12:107–117.!
81! !
Wilfong,!B.!N.,!D.!L.!Gorchov,!and!M.!C.!Henry.!2009.!Detecting!an!invasive!shrub!in! deciduous!forest!understories!using!remote!sensing.!Weed!Science!57:512–520.!! ' !
82! !
Table'1.!!!Browse!on!first?year!Lonicera%maackii)twigs!in!five!deer!access!plots!within! Miami!University!Natural!Areas.!! ' L.%maackii%! !Browsed! Percent! ' Site! Twigs!Sampled% !Twigs! Browsed! ' ! ' Bachelor!Preserve! 214! 12! 5.61! ' College!Woods! 148! 3! 2.03! ' Kramer!Preserve! 221! 12! 5.43! ' Reinhart!Preserve! 289! 5! 1.73! ' Western!Woods! 181! 3! 1.66! ' '
83! ! 300
250 * 200 150 100 50 Change in L. maackii BA /m^2) (cm^2 L. 200 in maackii Change 0
AC EX
Deer ! ' Figure'1.!Boxplot!of!Lonicera%maackii%basal!area!change!between!2010!and!2015.!!N!=!5! sites,!each!with!a!deer!access!(AC)!and!a!deer!exclosure!(EX)!plot.!!Asterisk!indicates! significant!difference!between!treatments!(P!
84! !
2 m
1.5 m
Deer Treatment Exclosure
Height Access
1 m *
0.5 m *
0 10 20 30 40 Lonicera maackii Frequency ! ' Figure'2.!Mean!frequency!of!Lonicera%maackii%foliage!at!each!of!four!heights!in!deer!access! and!deer!exclosure!plots.!!!Error!bars!are!SE!based!on!data!from!5!plots.!Asterisk!indicates! significant!difference!between!treatments!(P!
85! !
1 - 1.5 m *
Deer Treatment 0.5 - 1 m Exclosure * Access Height Range
0 - 0.5 m
0 20 40 60 Lonicera maackii Frequency ! ' Figure'3.!Mean!frequency!of!Lonicera%maackii%foliage!at!each!height!range!in!deer!access! and!deer!exclosure!plots.!!!Error!bars!are!SE!based!on!data!from!5!plots.!Asterisk!indicates! significant!difference!between!treatments!(P!
86! !
Chapter'5.'Conclusions'
Goals' This!dissertation!sought!to!investigate!the!interactions!between!an!overabundant,! generalist!herbivore,!white?tailed!deer,!and!a!common!invasive!shrub,!Lonicera%maackii.!I! found!that!interactions!between!deer!and!L.%maackii%on!tree!seedling!cover!are!non? additive!and!antagonistic!in!nature,!that!the!interactions!between!L.%maackii%and!native!tree! seedlings!are!facilitative,!and!that!deer!browse!on!L.%maackii%is!sufficient!to!alter!the!growth! and!structure!of!L.%maackii.!
Non:additive,'antagonistic'interactive'effects'on'plant'community'composition' In!Chapter!2,!I!investigated!the!interaction!between!these!two!factors!on!plant! species!richness!and!composition!to!address!the!hypothesis!that!while!both!L.%maackii%and! deer!individually!have!negative!impacts!on!forest!floor!plant!richness!and!composition,! when!taken!together,!those!impacts!will!be!non?additive!and!antagonistic!in!nature.!!In! order!to!do!so,!I!used!five!20m!x!20m!pairs!of!deer!exclosure!and!access!plots!within!the! Miami!University!Natural!Areas.!!Within!each!one!of!those!plots,!L.%maackii%was!removed! from!half!of!the!plot.!!I!utilized!this!split?plot!design!to!assess!plant!identity!and!cover,!and! draw!conclusions!from!the!crossed!effects.!!I!found!significant!effects!of!L.%maackii%removal! over!time!on!plant!species!richness,!but!minimal!effects!of!L.%maackii%on!plant!species! composition.!!I!found!significant!effects!of!L.%maackii%on!the!shrub,!tree,!annual,!spring! perennial!and!graminoid!functional!types;!each!of!these,!with!the!exception!of!shrub,!had! greater!cover!in!areas!of!L.%maackii%removal.!!I!did!not!find!an!effect!of!white?tailed!deer!on! species!richness,!but!did!find!very!significant!effects!of!deer!exclusion!on!spring!plant! species!composition!in!every!year!and!season!after!the!first.!!Indicators!of!deer!exclosure! included!Podophyllum%peltatum,!an!unpalatable!herb,%and!Acer%saccharum,%a!tree!species!of! moderate!deer!preference.!!I!found!an!effect!of!deer!on!cover!of!trees!in!the!forest!floor! layer,!but!not!on!cover!of!any!other!functional!types.!!I!found!an!interaction!between!L.% maackii%and!deer!on!tree!seedling!cover,!such!that!there!was!no!effect!of!deer!exclosure! when!L.%maackii%is!present.!!This!suggests!that!L.%maackii%mitigates!the!negative!effect!of! deer!browse!on!tree!seedling!cover,!and!facilitates!tree!seedling!survival!and!growth.!!This!
87! ! also!supports!the!hypothesis!that!the!relationship!between!deer!and!L.%maackii%is!non? additive!and!antagonistic!in!nature.!
Facilitative'interactions'between'L.)maackii)and'tree'seedlings) In!Chapter!3,!I!investigated!whether!L.%maackii%provides!a!refuge!for!tree!seedlings! from!browse!by!overabundant!white?tailed!deer,!testing!the!hypothesis!that!facilitation,! rather!than!competition,!was!the!dominant!interaction!between!L.%maackii!and!native!tree! species.!!This!study!utilized!ten,!large!focal!L.%maackii!shrubs!in!each!of!two!of!the!Miami! University!Natural!Areas,!assigning!half!to!be!fenced!deer!exclosures,!and!half!to!be!deer! access!plots.!!Native!tree!seedlings!were!planted!under!and!away!from!L.%maackii!in!areas! of!deer!access!and!exclosure.!!This!study!design!was!novel!in!investigating!whether!an! invasive!shrub!in!eastern!deciduous!forest!facilitates!tree!seedling!growth!and!survival.!!I! found!significant!interactions!of!deer!exclosure!and!position!under!or!away!from!L.%maackii% on!Acer%saccharum!survival!and!leaf!count,!which!suggested!that!location!under!the!canopy! of!L.%maackii%facilitates!the!survival!and!growth!of!tree!seedlings.!!I!found!minimal!evidence! for!competition!between!L.%maackii%and!tree!seedlings.!Therefore,!I!found!the!relationship! between!native!tree!seeding!and!L.%maackii!to!be!facilitative!rather!than!competitive!in! nature.!
Decreased'growth'and'cover'of'L.)maackii)in'the'presence'of'deer' ! In!Chapter!4,!!We!investigated!if!deer!browse!on!L.%maackii!is!low,!consistent!with! the!enemy!release!hypotheses!(ERH),!or!high,!consistent!with!the!biotic!resistance! hypothesis!(BRH),!and!!further,!if!this!browse!is!sufficient!to!alter!the!growth!and!structure! of!L.%maackii.%!Utilizing!the!L.%maackii!present!portion!of!the!large!deer!exclosures!and! controls!from!Chapter!2,!,!I!recorded!L.%maackii%leaf!frequency!between!0!and!2!m!from!the! ground.!!My!undergraduate!collaborator,!Christina!Haffey!also!measured!the!basal!area! (BA)!of!L.%maackii%in!each!plot,!and!measured!the!amount!of!browse!on!L.%maackii%shrubs% within!the!deer!access!plots.!!I!found!significantly!greater!basal!area!growth!between!2010! and!2015!in!the!deer!exclosure!plots.!!Within!the!deer!exclosures,!I!found!significantly! higher!frequency!of!L.%maackii%leaves!at!0.5!m,!between!0.5!and!1m,!at!1m!and!between!1! and!1.5!meters.!!These!differences!occurred!within!the!height!range!where!deer!browse,!
88! ! and!I!attributed!these!differences!in!leaf!frequency!to!deer!browse!in!the!deer!access!plots.!! These!findings!are!consistent!with!the!BRH,!in!that!a!native!species!could!suppress!an! invasive!plant!through!herbivory.!!!
Synthesis' The!facilitation!of!native!tree!seedlings!by!L.%maackii!is!both!novel!and!an!important! insight!into!the!interactions!between!L.%maackii%and!deer.!!Support!for!this!idea!is!not! limited!to!the!findings!from!Chapter!3,!as!the!interactive!effect!of!L.%maackii!and!deer!on! tree!functional!type!cover!(Chapter!2)!also!suggested!that!presence!L.%maackii%reduces!the! negative!effects!of!deer.!!I!would!argue!that!in!most!of!the!large!deer!access!plots!the! density!of!L.%maackii!shrubs!is!high!enough!within!L.%maackii%intact!portion!to!discourage! deer!from!navigating!through!them!or!function!as!a!continuous!physical!barrier!to!deer! herbivory.!!Additionally,!in!attempting!to!locate!large,!isolated!shrubs!to!implement!the! small!exclosure!experiment!(chapter!3),!I!determined!that!L.%maackii%shrubs!more! commonly!grew!in!these!dense!patches.!!I!suggest!then,!that!between!the!20!x!20!m!plots! and!the!isolated!large!shrub!experiment!my!investigations!covered!!!the!range!of!L.%maackii% densities!within!these!forests.!!I!conclude!that!location!directly!under!an!isolated!shrub!or! location!in!a!more!dense!stand!of!L.%maackii%can%function!to!protect!tree!seedlings!from! deer!browse.!! Non?additive,!antagonistic!interactions!can!come!about!in!two!ways!–!by!L.%maackii% mitigating!the!negative!effect!of!deer,!or!by!deer!mitigating!the!negative!effect!of!L.%maackii.!! The!significant!interactions!of!L.%maackii%and!deer!on!tree!cover!in!areas!of!deer!access! (Chapter!2)!suggest!that!the!presence!of!L.%maackii%discourages!deer!from!browsing!on! seedlings!in!areas!of!L.%maackii%presence,!likely!by!being!a!physical!barrier,!restricting!deer! access!to!seedlings.!!In!this!way,!L.%maackii%is!able!to!mitigate!the!negative!effect!of!deer!on! tree!seedlings.!However,!the!large!effect!of!deer!browse!on%L.%maackii!(Chapter!4)!supports! the!interpretation!of!deer!mitigation!of!the!negative!L.%maackii%effect!by!reducing!shrub! cover!and!canopy!size,!thus!lessening!the!negative!impacts!of!shading!by!L.%maackii!on! forest!floor!vegetation.!!In!either!case,!the!interaction!between!deer!and!L.%maackii!is!non? additive!and!the!combined!effects!of!L.%maackii!and!deer!are!less!negative!than!would!be! predicted!from!the!sum!of!their!effects.!!!!!
89! !
If!we!consider!the!facilitation!of!tree!seedlings!by!L.%maackii%in!light!of!the!finding!of! significant!deer!browse!on!L.%maackii,!as!well!as!the!suggestion!that!deer!may!have!only! recently!begun!recognizing!and!exploiting!L.%maackii%as!a!food!source,!we!are!left!with!a! particularly!damaging!scenario!for!tree!seedlings.!!!If!the!trend!of!deer!exploitation!of%L.% maackii!continues,!and!I!have!no!reason!to!believe!that!it!will!not,!it!is!likely!that!over!time! the!negative!effect!of!deer!on!L.%maackii%will!reduce!the!current!positive,!facilitative!effect!of! L.%maackii%on!tree!seedlings.!!This!effect!would!be!brought!about!through!deer!browse!on! the!branches!of!L.%maackii%less!than!2!m!from!the!ground,!the!top!of!the!height!range!of!deer! browse.!!As!browsing!eliminates!those!branches,!the!arching!canopy!that!protects!the!tree! seedlings!from!browse!is!lost,!leading!to!an!even!larger!net!negative!effect!of!deer!on!tree! seedlings.!!! For!these!reasons,!I!suggest!that!if!land!and!forest!managers!have!a!goal!of!increased!tree! regeneration,!they!should!focus!their!limited!resources!on!the!reduction!of!deer! populations,!rather!than!the!removal!of!L.%maackii.%While!deer!herds!are!large!and! exhibiting!high!browse!pressure!in!forests,!the!current!relationship!between!L.%maackii%and! tree!seedlings!is!facilitative,!protecting!seedlings!from!the!impacts!of!these!deer.!!If!current! deer!populations!are!sustained!or!if!they!increase,!greater!deer!browse!on!L.%maackii!will! continue!to!increase,!lessening!their!ability!to!protect!seedlings.!!Once!deer!herds!are! brought!back!down!to!lower!densities,!the!status!of!L.%maackii%as!facilitative!to!tree!growth! and!survival!should!be!reassessed.!!Facilitative!plant?to?plant!interactions!are!only! predicted!in!stressful!environments,!such!as!one!prompted!by!heavy!deer!browse,!and!if! this!browse!pressure!is!alleviated,!the!interaction!between!L.%maackii!and!tree!seedlings! would!change!to!one!that!is!competitive!in!nature.!!!Management!funds!could,!at!that!point,! be!redirected!to!removal!of!L.%maackii.!!!!While!these!studies!specifically!addressed!the! interactions!between!deer!and!L.%maackii,!one!could!speculate!that!where!deer!pressure!is! sufficiently!high,!any!shrub!that!deer!did!not!prefer!as!browse!and/or!that!was!heavily! defended!by!thorns!or!spines!should!function!to!protect!tree!seedlings!from!deer.!!If!this! facilitation!outweighed!any!competitive!effects!of!the!shrub!on!tree!seedlings,!the!seedlings! should!be!able!to!grow!to!the!point!where!they!overtop!their!facilitator!shrub,!and!can!then! recruit!into!the!canopy,!unharmed!by!deer!browse.! !
90! !
Appendix''
Additional'Tables'and'Figures'from'Chapter'2' ! ' Appendix'Table'A1.!Sampling!dates.!! Year! Spring! Summer! ! ! 2011! June!1!?!14! July!12!?!18! ! 2012! May!6!?!10! June!24!?!27! ! 2013! May!20!?!24! July!9!?!14! ! 2014! June!12?!17! July!16!?!22! ! ! ' ' '
91! !
Appendix'Table'A2.'Species!cover!(mean!and!standard!error)!by!treatment!type.'' ' Spring'2011'
Mean$Cover$in$$cm^2/1800$cm^2$ Standard$Error Deer$Exclosure Deer$Exclosure Deer$Access Deer$Access Deer$Exclosure Deer$Exclosure Deer$Access Deer$Access L.#maackii#Removed L.#maackii#Intact L.#maackii#Removed L.#maackii#Intact L.#maackii#Removed L.#maackii#Intact L.#maackii#Removed L.#maackii#Intact Acer#saccharum 836 853 1264 998 448 425 516 344 Aesculus#sp.# 149 410 101 541 95 155 48 182 Ageratina#altissima 1407 1023 982 680 1214 413 361 257 Alliaria#petiolata 239 290 264 299 78 94 100 163 Allium#tricoccum 226 341 702 128 84 161 234 38 Arisaema#triphyllum 30 169 275 129 24 50 101 79 Botrychium#virginianus 476 291 218 286 351 148 104 75 Cardamine#douglassii 122 168 540 429 98 98 360 206 Carex#aurea 259 196 115 70 173 170 79 49 Carpinus#caroliniana 245 378 80 108 175 212 58 108 Carya#sp.# 122 111 42 47 122 68 18 31 Celtis#occidentalis 130 78 170 39 130 55 170 27 Cercis#canadensis 71 66 41 20 46 66 23 20 Delphinium#tricorne 6 66 28 91 6 47 28 27 Dicentra#cucullaria 269 62 4 43 71 62 4 15 Euonymus#fortunei 17 8 57 27 17 5 43 21 Fragaria#vesca 0 51 45 66 0 51 45 57 Fraxinus#sp 132 12 12 20 46 12 12 20 Galium#aparine 38 27 58 35 38 17 36 20 Galium#circaezans 326 260 80 0 260 147 80 0 Galium#mollugo 11 7 10 0 7 7 10 0 Geranium#maculatum 45 6 12 123 28 6 12 59 Glechoma#hederacea 36 36 14 16 19 29 14 16 Hydrophyllum#macrophyllum 0 0 165 6 0 0 165 6 Ilex#opaca 33 56 15 40 24 49 10 24 Impatiens#sp.# 4 0 62 22 4 0 62 14 Leersia#sp.# 0 128 6 2 0 128 6 2 Ligustrum#vulgare 2 29 10 8 2 23 6 8 Liriodendron#tulipfera 30 0 86 15 30 0 53 12 Lonicera#japonica 0 47 48 60 0 47 48 60 Lonicera#maackii 2 39 0 81 2 25 0 68 Ophioglossum#pusillum 14 20 42 0 14 20 21 0 Oxalis#sp.# 0 0 66 114 0 0 66 114 Parthenocissus#quinquefolia 0 8 25 16 0 8 13 16 Pilea#pumilla 38 0 39 0 25 0 30 0 Poa#sp.# 0 24 112 8 0 24 73 8 Podophyllum#peltatum 7 0 11 0 7 0 11 0 Polyogonum#sp.# 18 44 0 6 18 31 0 6 Polyongatum#biflorum 0 0 32 93 0 0 32 93 Quercus#sp.# 16 0 0 0 10 0 0 0 Ribes#cynosbati# 12 0 2 4 12 0 2 4 Robinia#pseudoacacia 0 24 20 15 0 24 20 15 Rosa#multiflora 0 0 18 4 0 0 11 4 Sanguinaria#canadensis 16 0 0 0 16 0 0 0 Sanicula#canadensis 54 0 5 0 54 0 5 0 Sanicula#odorata 0 0 0 38 0 0 0 38 Sassafras#albidum 0 0 100 0 0 0 100 0 Smilax#sp.# 36 8 0 0 36 8 0 0 Stellaria#media 15 0 0 0 15 0 0 0 Tilia#americana 50 0 20 0 50 0 20 0 Toxicodendron#radicans 2 0 0 6 2 0 0 6 Tradescantia#sp 0 0 0 13 0 0 0 13 Trillium#sp.# 0 0 18 4 0 0 18 4 Viola#sp 12 0 0 0 12 0 0 0 Vitis#sp.# 0 0 40 0 0 0 40 0 Unknown$Species$SP2011F1g 0 5 16 0 0 5 16 0 Unknown$Species$SP2011F2 0 0 6 75 0 0 6 75 Unknown$Species$SP2011F3 26 6 0 0 26 6 0 0 Unknown$Species$SP2011F4 20 100 0 0 20 100 0 0 Unknown$Species$SP2011F5g 36 0 0 0 36 0 0 0 Unknown$Species$SP2011F6 6 0 0 0 6 0 0 0 Unknown$Species$SP2011F7g 0 12 0 0 0 12 0 0 Unknown$Species$SP2011F8 0 0 0 100 0 0 0 100 Unknown$Species$SP2011F9 0 0 0 8 0 0 0 8 Unknown$Species$SP2011F10 0 6 0 0 0 6 0 0 Unknown$Species$SP2011F11 0 0 60 0 0 0 60 0 Unknown$Species$SP2011F12 0 0 0 10 0 0 0 10 Unknown$Species$SP2011F13 6 0 0 0 6 0 0 0 Unknown$Species$SP2011F14 4 0 0 0 4 0 0 0 Unknown$Species$SP2011F15 0 20 0 0 0 20 0 0 Unknown$Species$SP2011F16 0 0 0 15 0 0 0 15 Unknown$Species$SP2011F17 0 0 25 0 0 0 25 0 Unknown$Species$SP2011F18 2 0 0 0 2 0 0 0 *$g$after$the$unknown$species$number$refers$to$a$graminoidFlike$species '
92! !
Spring'2012' Mean$Cover$in$$cm^2/1800$cm^2$ Standard$Error Deer$Exclosure Deer$Exclosure Deer$Access Deer$Access Deer$Exclosure Deer$Exclosure Deer$Access Deer$Access L.#maackii#Removed L.#maackii#Intact L.#maackii#Removed L.#maackii#Intact L.#maackii#Removed L.#maackii#Intact L.#maackii#Removed L.#maackii#Intact Acer#rubrum 0 0 4 0 0 0 4 0 Acer#saccharum 203 117 114 145 81 81 50 50 Aesculus#sp.# 60 0 60 0 60 0 60 0 Alliaria#petiolata 198 194 136 194 108 80 73 122 Allium#sp. 0 0 16 4 0 0 16 4 Allium#tricoccum 1 0 1 2 1 0 1 2 Andropogon#gerardi 20 0 12 30 20 0 12 30 Arisaema#triphyllum 0 5 0 5 0 5 0 5 Berberis#thunbergii 12 0 0 0 12 0 0 0 Boehmeria#cylindrica 152 225 393 64 105 131 155 34 Botrychium#virginianus 14 0 154 9 14 0 80 6 Cardamine#concatenata 6 0 32 0 5 0 10 0 Cardamine#douglassii 60 0 0 40 60 0 0 40 Carpinus#caroliniana 12 0 0 0 12 0 0 0 Carya#sp. 84 30 0 18 59 13 0 18 Cercis#canadensis 10 2 0 6 10 2 0 6 Claytonia#virginica 5 13 38 9 3 13 23 7 Erythronium#americanum 0 0 2 0 0 0 2 0 Euonymus#alatus 0 18 42 26 0 18 42 26 Euonymus#fortunei 10 0 0 5 10 0 0 5 Fragaria#vesca 40 31 45 97 40 31 45 78 Fraxinus#sp.# 148 128 116 105 61 68 80 42 Galium#aparine 194 322 188 126 79 137 116 55 Galium#circaezans 24 6 19 18 16 6 13 12 Galium#mollugo 8 0 86 26 8 0 40 26 Geum#canadense 56 8 0 41 56 8 0 41 Glechoma#hederacea 0 0 16 0 0 0 15 0 Hydrophyllum#macrophyllum 0 0 40 34 0 0 40 16 Impatiens#sp.# 10 110 178 329 6 49 139 154 Ligustrum#vulgare 77 192 32 100 58 192 32 71 Liriodendron#tulipfera 0 0 1 0 0 0 1 0 Lonicera#japonica 20 0 0 0 20 0 0 0 Lonicera#maackii 98 290 84 330 68 149 51 134 Lonicera#morrowii 0 0 0 24 0 0 0 24 Ophioglossum#pusillum 33 11 2 0 26 11 2 0 Ostrya#virginiana 13 0 0 0 12 0 0 0 Oxalis#sp.# 0 0 0 1 0 0 0 1 Parthenocissus#quinquefolia 101 99 347 217 69 65 114 95 Poa#sp.# 166 180 159 154 114 100 100 83 Podophyllum#peltatum 750 480 182 0 325 227 92 0 Polyongatum#biflorum 203 120 54 116 59 64 26 61 Prunus#serotina 57 152 100 63 53 68 81 38 Quercus#sp. 20 0 0 0 13 0 0 0 Ranunculus#abortivus 0 0 4 0 0 0 4 0 Rhamnus#cathartica 0 0 0 6 0 0 0 6 Ribes#cynosbati# 0 0 40 2 0 0 40 2 Rosa#multiflora 0 20 0 0 0 20 0 0 Sanguinaria#canadensis 45 27 58 6 45 25 51 6 Sanicula#odorata 1753 1134 781 637 1403 468 230 178 Scutellaria#sp. 25 0 0 15 25 0 0 15 Smilax#sp.# 6 42 6 2 6 22 6 2 Stellaria#media 382 520 985 375 377 402 942 353 Stellaria#pubera 89 0 30 12 55 0 30 12 Thalictrum#thalictroides 15 14 0 0 15 14 0 0 Toxenodendron#radicans 12 36 0 0 12 36 0 0 Trillium#sessile 26 0 0 20 26 0 0 20 Trillum#sp.# 0 0 22 66 0 0 16 49 Ulmus#rubra 26 16 10 20 19 16 6 20 Viola#sp. 35 136 332 151 22 70 160 91 Vitis#sp.# 0 19 4 20 0 18 4 13 Unknown$Species$SP2012F1 0 0 110 20 0 0 68 20 Unknown$Species$SP2012F2 0 0 6 0 0 0 6 0 Unknown$Species$SP2012F3 0 0 10 40 0 0 10 40 Unknown$Species$SP2012F4 60 0 0 20 60 0 0 20 Unknown$Species$SP2012F5 40 0 0 0 40 0 0 0 Unknown$Species$SP2012F6 20 0 12 12 20 0 12 12 Unknown$Species$SP2012F7 0 5 0 0 0 5 0 0 Unknown$Species$SP2012F8 0 0 12 0 0 0 12 0 Unknown$Species$SP2012F9 0 0 25 0 0 0 25 0 Unknown$Species$SP2012F10 0 0 60 0 0 0 60 0 Unknown$Species$SP2012F11 0 0 0 60 0 0 0 60 Unknown$Species$SP2012F12g 60 52 40 0 60 52 30 0 Unknown$Species$SP2012F13g 0 0 24 0 0 0 24 0 Unknown$Species$SP2012F14g 88 34 77 40 48 27 49 18 *$g$after$the$unknown$species$number$refers$to$a$graminoidFlike$species '
93! !
Spring'2013'
Mean$Cover$in$$cm^2/1800$cm^2$ Standard$Error Deer$Exclosure Deer$Exclosure Deer$Access Deer$Access Deer$Exclosure Deer$Exclosure Deer$Access Deer$Access L.#maackii#Removed L.#maackii#Intact L.#maackii#Removed L.#maackii#Intact L.#maackii#Removed L.#maackii#Intact L.#maackii#Removed L.#maackii#Intact Acer#saccharum 147 59 76 97 74 34 46 56 Aesculus#sp.# 40 0 0 40 40 0 0 40 Alliaria#petiolata 113 91 396 200 98 89 374 177 Allium#tricoccum 1 1 2 0 1 1 2 0 Arisaema#triphyllum 0 4 49 32 0 4 43 32 Asarum#canadense 0 0 60 36 0 0 60 36 Boehmeria#cylindrica 163 88 282 168 157 68 102 104 Botrychium#virginianus 25 2 1 0 25 2 1 0 Cardamine#concatenata 4 20 32 7 3 18 15 7 Cardamine#douglassii 50 0 0 0 50 0 0 0 Carya#sp. 12 12 0 60 12 12 0 38 Cercis#canadensis 2 8 0 0 2 6 0 0 Claytonia#virginica 1 2 2 1 1 2 1 1 Dicentra#sp.# 39 0 17 0 39 0 17 0 Euonymus#alata 50 0 0 0 50 0 0 0 Euonymus#fortunei 46 0 0 0 41 0 0 0 Fagus#grandifolia# 0 0 0 40 0 0 0 40 Festuca#sp. 23 5 34 28 14 5 22 19 Fraxinus#sp.# 291 39 51 24 140 27 21 12 Galium#aparine 120 15 124 44 74 13 41 7 Galium#circaezans 4 0 0 0 2 0 0 0 Galium#mollugo 18 6 0 5 18 6 0 5 Glechoma#hederacea 0 0 8 0 0 0 8 0 Hydrophyllum#virginianum 6 0 0 148 6 0 0 141 Impatiens#sp.# 13 15 22 58 13 15 16 36 Leersia#sp 103 81 153 27 36 42 65 14 Ligustrum#vulgare 70 276 5 86 70 169 5 86 Lonicera#japonica 14 6 0 0 14 6 0 0 Lonicera#maackii 73 165 136 449 39 116 84 245 Lonicera#morrowii 0 0 0 12 0 0 0 12 Ostrya#virginiana 20 0 0 20 20 0 0 20 Oxalis#sp.# 4 0 0 0 4 0 0 0 Parthenocissus#quinquefolia 77 199 235 78 42 110 133 51 Podophyllum#peltatum 443 382 302 0 178 193 195 0 Polygonum#sp 0 0 11 6 0 0 11 6 Polyongatum#biflorum 260 59 75 75 104 49 40 28 Prunus#serotina 139 74 71 48 111 34 51 18 Ribes#cynosbati# 0 40 20 0 0 40 20 0 Rosa#multiflora 0 20 0 0 0 20 0 0 Sanguinaria#canadensis 0 25 0 0 0 25 0 0 Sanicula#canadensis 37 12 30 100 25 12 30 100 Sanicula#odorata 1023 844 398 303 848 424 190 152 Sassafras#albidum 2 0 0 0 2 0 0 0 Smilax#sp.# 42 12 32 16 42 12 21 10 Stellaria#media 89 15 157 30 89 15 157 30 Stellaria#pubera 28 6 0 0 28 6 0 0 Thalictrum#thalictroides 5 0 0 0 3 0 0 0 Toxenodendron#radicans 14 0 0 0 14 0 0 0 Trifolium#sp 0 0 2 0 0 0 2 0 Trillium#sp.# 12 45 11 0 12 45 11 0 Ulmus#rubra 34 12 2 0 22 12 2 0 Viola#sp. 161 20 168 24 100 20 87 12 Vitis#sp.# 14 0 5 16 7 0 5 16 Unknown$Species$SP2013F1 1 0 3 0 1 0 3 0 Unknown$Species$SP2013F2g 0 0 12 0 0 0 12 0 Unknown$Species$SP2013F3g 0 20 108 30 0 20 47 20 Unknown$Species$SP2013F4 36 30 35 0 36 30 35 0 Unknown$Species$SP2013F5 0 0 35 0 0 0 35 0 Unknown$Species$SP2013F6 60 0 0 0 60 0 0 0 Unknown$Species$SP2013F7g 0 0 12 2 0 0 12 2 Unknown$Species$SP2013F8 0 0 0 12 0 0 0 12 Unknown$Species$SP2013F9 0 0 0 40 0 0 0 40 *$g$after$the$unknown$species$number$refers$to$a$graminoidFlike$species '
94! !
Spring'2014
Mean$Cover$in$$cm^2/1800$cm^2$ Standard$Error Deer$Exclosure Deer$Exclosure Deer$Access Deer$Access Deer$Exclosure Deer$Exclosure Deer$Access Deer$Access L.#maackii#Removed L.#maackii#Intact L.#maackii#Removed L.#maackii#Intact L.#maackii#Removed L.#maackii#Intact L.#maackii#Removed L.#maackii#Intact Acer#saccharum 282 155 94 56 40 56 36 23 Alliaria#petiolata 203 187 406 252 142 88 204 132 Allium#tricoccum 12 12 4 9 12 11 4 9 Arisaema#triphyllum 0 0 23 12 0 0 15 12 Asarum#canadense 0 0 72 0 0 0 72 0 Boehmeria#cylindrica 276 125 298 45 233 79 184 22 Botrychium#virginianum 58 12 18 0 58 10 14 0 Cardamine#concatenata 4 0 0 0 4 0 0 0 Cardamine#douglassii 73 0 0 0 45 0 0 0 Carex#aurea# 8 20 0 6 8 20 0 4 Carpinus#caroliniana 0 0 0 50 0 0 0 32 Carya#sp. 24 10 0 0 15 10 0 0 Celastrus#orbiculatus 0 0 36 68 0 0 29 68 Cercis#canadensis 16 5 0 0 10 3 0 0 Circaea#lutetiana 0 0 40 106 0 0 40 106 Cornus#florida 0 0 40 0 0 0 40 0 Cystopteris#fragilis 0 0 28 0 0 0 28 0 Elymus#sp. 30 0 0 0 30 0 0 0 Euonymus#alata 90 40 0 0 60 40 0 0 Euonymus#fortunei 76 8 0 16 69 8 0 16 Fragaria#vesca 2 0 0 0 2 0 0 0 Fraxinus#quadrangulata 84 0 0 0 84 0 0 0 Fraxinus#sp.# 347 143 266 141 171 66 87 38 Galium#aparine 16 0 67 10 16 0 51 10 Galium#mollugo 5 0 4 8 5 0 4 6 Geranium#maculatum 0 25 0 0 0 25 0 0 Glechoma#hederacea 0 0 40 0 0 0 40 0 Hydrophyllum#macrophyllum 0 0 0 24 0 0 0 24 Hystrix#patula 0 0 16 0 0 0 10 0 Impatiens#sp.# 6 104 36 72 6 64 36 51 Leersia#sp. 75 19 46 42 27 13 20 25 Ligustrum#vulgare 2 240 60 144 2 154 60 99 Liriodendron#tulipfera 2 0 4 0 2 0 4 0 Lonicera#japonica 188 24 0 0 162 24 0 0 Lonicera#maackii 153 418 152 299 82 173 89 110 Ophioglossum#pusillum 13 11 3 4 6 11 3 4 Ostrya#virginiana 0 0 0 12 0 0 0 12 Parthenocissus#quinquefolia 278 62 370 268 137 39 205 117 Pilea#pumila 0 0 0 59 0 0 0 46 Podophyllum#peltatum 673 372 257 0 465 243 156 0 Polygonum#sp. 0 0 0 62 0 0 0 50 Polyongatum#biflorum 142 14 82 46 76 12 42 34 Prunus#serotina 178 225 48 88 161 194 33 77 Quercus#sp.# 0 0 18 0 0 0 18 0 Ribes#cynosbati# 0 0 0 6 0 0 0 6 Rosa#multiflora 0 50 6 0 0 50 6 0 Sanguinaria#canadensis 25 0 0 0 25 0 0 0 Sanicula#canadensis 30 0 0 0 30 0 0 0 Sanicula#odorata 995 756 312 172 870 463 131 83 Setaria#lutescens 0 0 78 18 0 0 60 18 Smilax#sp.# 102 52 24 0 65 39 17 0 Stellaria#media 7 4 27 0 7 4 26 0 Thalictrum#thalictroides 10 1 0 0 10 1 0 0 Toxicodendron#radicans 16 5 63 0 9 5 39 0 Trillium#sp.# 0 12 0 39 0 12 0 25 Ulmus#rubra 90 25 20 20 46 19 11 13 Viola#sp. 109 46 178 37 81 23 130 24 Vitis#sp.# 39 30 40 30 14 10 28 14 Unknown$Species$SP2014F1 40 20 0 0 40 20 0 0 Unknown$Species$SP2014F2 40 0 4 3 28 0 4 3 Unknown$Species$SP2014F3g 0 10 70 56 0 10 70 56 Unknown$Species$SP2014F4 50 6 6 12 50 6 6 12 Unknown$Species$SP2014F5 0 0 3 0 0 0 3 0 Unknown$Species$SP2014F6 0 0 0 6 0 0 0 6 Unknown$Species$SP2014F7 120 0 0 0 120 0 0 0 Unknown$Species$SP2014F8g 40 0 0 6 40 0 0 6 Unknown$Species$SP2014F9g 0 0 102 0 0 0 65 0 *$g$after$the$unknown$species$number$refers$to$a$graminoidFlike$species '
95! !
Summer'2011' Mean$Cover$in$$cm^2/1800$cm^2$ Standard$Error Deer$Exclosure Deer$Exclosure Deer$Access Deer$Access Deer$Exclosure Deer$Exclosure Deer$Access Deer$Access L.#maackii#Removed L.#maackii#Intact L.#maackii#Removed L.#maackii#Intact L.#maackii#Removed L.#maackii#Intact L.#maackii#Removed L.#maackii#Intact Acer#saccharum# 264 169 177 126 85 86 111 39 Ageratina#altissima 0 0 40 6 0 0 40 6 Alliaria#petiolata 525 426 423 435 270 209 280 190 Allium#tricoccum 4 14 13 2 4 14 10 2 Arisaema#triphyllum 24 20 72 40 24 20 72 30 Botrychium#virginianus 9 16 42 41 9 12 42 26 Cardamine#douglassii 6 0 0 0 6 0 0 0 Carya#sp.# 141 52 90 59 113 39 63 28 Celtis#occidentalis 24 0 0 12 19 0 0 12 Cercis#canadensis 58 56 63 28 51 50 50 28 Elymus#sp.# 24 15 0 0 24 15 0 0 Elymus#hystrix 0 0 0 0 0 0 0 0 Euonymus#fortunei 11 0 0 14 11 0 0 9 Festuca#sp. 126 84 6 39 126 84 6 39 Fragaria#vesca 14 0 0 0 12 0 0 0 Fraxinus#sp.# 136 185 214 182 40 56 120 71 Galium#aparine 20 5 49 0 20 5 49 0 Galium#circaezans 104 3 16 18 55 2 10 14 Galium#mollugo 349 222 189 358 251 163 170 358 Glechoma#hederacea 5 0 0 0 5 0 0 0 Impatiens#sp.# 501 104 325 205 328 64 222 150 Leersia#sp.# 163 108 157 102 98 79 66 52 Ligustrum#vulgare 10 513 166 8 10 211 91 8 Liriodendron#tulipfera 33 14 0 52 22 10 0 52 Lonicera#japonica 12 0 0 36 12 0 0 36 Lonicera#maackii 234 71 171 266 168 30 88 116 Ophioglossum#pusillum 12 0 0 0 12 0 0 0 Parthenocissus#quinquefolia 306 224 316 218 103 104 119 144 Pilea#pumilla 16 165 357 198 10 93 186 105 Podophyllum#peltatum 60 0 0 0 60 0 0 0 Polygonum#sp. 0 6 110 96 0 6 68 82 Polyongatum#biflorum 235 90 227 31 168 68 220 16 Prunus#serotina 58 13 8 76 53 13 8 76 Sanguinaria#canadensis 8 0 0 40 8 0 0 40 Sanicula#odorata 823 640 738 534 574 268 297 239 Smilax#sp.# 31 32 25 6 24 32 25 6 Solanum#pumilum# 61 0 0 0 38 0 0 0 Taraxacum#officinale 0 0 18 0 0 0 18 0 Toxicodendron#radicans 38 37 0 4 38 37 0 4 Viola#sp. 118 16 165 10 80 10 75 7 Vitis#sp. 118 7 12 0 104 4 12 0 Unknown$Species$SUM2011F1g 26 56 2 0 21 49 2 0 Unknown$Species$SUM2011F2g 96 60 37 18 58 60 23 18 Unknown$Species$SUM2011F3g 0 0 159 16 0 0 107 16 Unknown$Species$SUM2011F4 73 132 12 71 53 118 12 47 Unknown$Species$SUM2011F5 40 0 42 28 40 0 42 28 Unknown$Species$SUM2011F6 0 28 0 16 0 28 0 16 Unknown$Species$SUM2011F7 0 0 26 12 0 0 16 12 Unknown$Species$SUM2011F8 60 0 0 9 60 0 0 9 Unknown$Species$SUM2011F9 0 0 0 95 0 0 0 95 Unknown$Species$SUM2011F10 0 8 0 0 0 8 0 0 Unknown$Species$SUM2011F11 0 0 5 0 0 0 5 0 Unknown$Species$SUM2011F12 0 0 0 9 0 0 0 9 Unknown$Species$SUM2011F13 0 15 0 0 0 15 0 0 *$g$after$the$unknown$species$number$refers$to$a$graminoidFlike$species '
96! !
Summer'2012'
Mean$Cover$in$$cm^2/1800$cm^2$ Standard$Error Deer$Exclosure Deer$Exclosure Deer$Access Deer$Access Deer$Exclosure Deer$Exclosure Deer$Access Deer$Access L.#maackii#Removed L.#maackii#Intact L.#maackii#Removed L.#maackii#Intact L.#maackii#Removed L.#maackii#Intact L.#maackii#Removed L.#maackii#Intact Acer#rubrum 0 0 0 12 0 0 0 12 Acer#saccharum 191 134 59 80 63 73 46 51 Alliaria#petiolata 99 49 20 33 42 27 13 14 Allium#tricoccum 12 6 11 8 12 6 7 8 Arisaema#triphyllum 0 0 8 12 0 0 8 12 Boehmeria#cylindrica 314 122 514 257 177 104 157 88 Botrychium#virginianus 24 0 28 0 24 0 14 0 Cardamine#douglassii 82 6 0 36 51 6 0 36 Carya#sp. 183 37 25 0 68 23 25 0 Cercis#canadensis 6 8 0 4 6 8 0 4 Elymus#hystrix 0 0 17 6 0 0 17 6 Elymus#sp. 90 34 66 20 90 24 35 13 Euonymus#fortunei 52 0 0 0 47 0 0 0 Festuca#sp. 142 127 62 31 119 113 41 31 Fragaria#vesca 0 0 12 0 0 0 12 0 Fraxinus#sp.# 224 51 88 25 163 26 44 12 Galium#aparine 48 18 66 32 48 18 42 23 Galium#circaezans 36 0 13 37 23 0 13 22 Galium#mollugo 0 0 86 60 0 0 56 50 Geum#canadense 80 40 65 0 80 40 65 0 Glechoma#hederacea 15 0 28 0 15 0 28 0 Impatiens#sp.# 12 19 16 83 12 19 15 36 Leersia#sp.# 0 0 0 0 0 0 0 0 Ligustrum#vulgare 66 171 26 70 59 105 19 56 Lonicera#maackii 38 158 164 154 20 87 93 86 Oxalis#sp.# 0 0 0 5 0 0 0 5 Parthenocissus#quinquefolia 160 80 348 165 39 36 144 97 Poa#sp.# 13 18 130 34 12 12 80 27 Podophyllum#peltatum 0 0 50 0 0 0 50 0 Polygonum#sp. 25 0 74 16 25 0 64 16 Polyongatum#biflorum 66 92 7 37 48 45 4 25 Prunus#serotina 128 82 79 81 100 60 50 48 Quercus#sp. 8 0 0 0 8 0 0 0 Ribes#cynosbati# 0 40 6 0 0 40 6 0 Rosa#multiflora 0 26 0 6 0 26 0 6 Sanguinaria#canadensis 40 0 80 0 40 0 80 0 Sanicula#odorata 1302 969 762 568 905 455 241 135 Smilax#sp.# 24 0 32 24 15 0 23 24 Toxenodendron#radicans 0 0 20 6 0 0 20 6 Ulmus#rubra 25 67 7 23 19 41 5 17 Viola#sp. 36 88 180 0 24 83 86 0 Vitis#sp. 31 2 13 12 13 2 7 12 Unknown$Species$SUM2012F1 40 0 0 5 40 0 0 5 Unknown$Species$SUM2012F2g 34 36 0 0 34 36 0 0 Unknown$Species$SUM2012F3g 0 0 14 0 0 0 14 0 Unknown$Species$SUM2012F4 48 0 0 0 48 0 0 0 Unknown$Species$SUM2012F5 0 0 0 44 0 0 0 44 Unknown$Species$SUM2012F6 0 0 12 0 0 0 12 0 Unknown$Species$SUM2012F7 20 0 0 0 20 0 0 0 Unknown$Species$SUM2012F8g 36 80 52 106 36 80 39 99 Unknown$Species$SUM2012F9 0 60 0 20 0 60 0 20 *$g$after$the$unknown$species$number$refers$to$a$graminoidFlike$species '
97! !
Summer'2013
Mean$Cover$in$$cm^2/1800$cm^2$ Standard$Error Deer$Exclosure Deer$Exclosure Deer$Access Deer$Access Deer$Exclosure Deer$Exclosure Deer$Access Deer$Access L.#maackii#Removed L.#maackii#Intact L.#maackii#Removed L.#maackii#Intact L.#maackii#Removed L.#maackii#Intact L.#maackii#Removed L.#maackii#Intact Acer#negundo 0 0 0 20 0 0 0 20 Acer#rubrum 6 0 0 0 4 0 0 0 Acer#saccharum 179 77 45 2 86 26 20 2 Aesculus#sp.# 48 0 0 0 48 0 0 0 Alliaria#petiolata 53 43 88 35 39 27 61 21 Allium#tricoccum 3 3 5 5 3 3 5 3 Arisaema#triphyllum 2 0 11 80 2 0 7 51 Asarum#canadense 0 0 35 0 0 0 35 0 Boehmeria#cylindrica 364 65 260 152 320 42 111 72 Botrychium#virginianus 12 0 0 0 12 0 0 0 Cardamine#douglassii 42 0 0 0 27 0 0 0 Carpinus#caroliniana 0 0 6 0 0 0 6 0 Carya#sp.# 17 0 0 0 15 0 0 0 Celtis#occidentalis 0 0 0 60 0 0 0 60 Cercis#canadensis 15 5 0 5 14 5 0 5 Circaea#lutetiana 60 10 62 188 60 10 50 188 Elymus#hystrix 0 6 82 12 0 6 53 12 Euonymus#alata 300 0 2 0 148 0 2 0 Euonymus#fortunei 54 0 0 5 54 0 0 5 Fagus#grandifolia# 0 0 0 50 0 0 0 50 Fraxinus#sp.# 284 66 41 25 121 56 26 14 Galium#aparine 18 6 76 0 18 6 76 0 Galium#circaezans 32 0 11 2 16 0 9 2 Galium#mollugo 6 2 11 38 6 2 11 23 Geranium#maculatum 0 6 0 0 0 6 0 0 Geum#canadense 0 16 0 2 0 16 0 2 Glechoma#hederacea 1 0 46 0 1 0 46 0 Hydrophyllum#virginianum 0 0 0 15 0 0 0 15 Impatiens#sp.# 16 16 121 60 10 16 96 54 Leersia#sp. 81 40 47 73 35 29 29 35 Ligustrum#vulgare 80 208 86 80 80 145 79 80 Liriodendron#tulipfera 0 0 0 1 0 0 0 1 Lonicera#japonica 67 30 0 0 41 30 0 0 Lonicera#maackii 132 259 133 618 91 138 65 210 Oxalis#sp.# 0 0 0 5 0 0 0 5 Parthenocissus#quinquefolia 163 12 246 110 62 12 107 46 Podophyllum#peltatum 60 0 60 0 60 0 60 0 Polygonum#sp. 0 0 26 6 0 0 26 4 Polyongatum#biflorum 150 0 9 0 56 0 9 0 Prunus#serotina 120 119 82 243 94 84 50 178 Ribes#cynosbati# 0 40 0 0 0 40 0 0 Rosa#multiflora 0 30 0 0 0 30 0 0 Sanguinaria#canadensis 20 0 0 0 20 0 0 0 Sanicula#canadensis 14 41 18 62 14 41 18 48 Sanicula#odorata 1157 849 281 142 1033 553 136 66 Smilax#sp.# 97 15 23 12 60 15 14 12 Toxenodendron#radicans 5 6 0 0 5 6 0 0 Ulmus#rubra 53 4 21 0 30 2 16 0 Viola#sp. 137 15 79 0 84 9 79 0 Vitis#sp.# 20 10 62 12 12 8 24 7 Unknown$Species$SUM2013F1g 22 64 84 28 14 41 30 23 Unknown$Species$SUM2013F2 60 4 0 0 60 4 0 0 Unknown$Species$SUM2013F3 0 30 0 0 0 30 0 0 Unknown$Species$SUM2013F4 0 0 18 0 0 0 18 0 Unknown$Species$SUM2013F5 0 0 18 0 0 0 18 0 Unknown$Species$SUM2013F6g 12 0 0 0 12 0 0 0 *$g$after$the$unknown$species$number$refers$to$a$graminoidFlike$species '
98! !
Summer'2014'
Mean$Cover$in$$cm^2/1800$cm^2$ Standard$Error Deer$Exclosure Deer$Exclosure Deer$Access Deer$Access Deer$Exclosure Deer$Exclosure Deer$Access Deer$Access L.#maackii#Removed L.#maackii#Intact L.#maackii#Removed L.#maackii#Intact L.#maackii#Removed L.#maackii#Intact L.#maackii#Removed L.#maackii#Intact Acer#saccahrum 372 276 58 118 43 102 27 50 Alliaria#petiolata 115 77 246 215 79 38 79 95 Allium#tricoccum 8 2 11 3 8 2 11 3 Arisaema#triphyllum 12 30 92 14 12 30 78 12 Asarum#canadense 0 0 54 80 0 0 54 80 Boehmeria#cylindrica 110 52 230 131 110 52 120 89 Botrychium#virginianum 38 0 0 0 38 0 0 0 Cardamine#douglassii 0 0 1 0 0 0 1 0 Carex#aurea# 50 0 6 0 50 0 6 0 Carya#sp. 44 24 0 0 31 16 0 0 Celastrus#orbiculatus 0 0 0 12 0 0 0 12 Cystopteris#fragilis 0 0 20 0 0 0 20 0 Dicentra#cucullaria 0 0 2 0 0 0 2 0 Elymus#sp. 12 0 0 0 12 0 0 0 Elymus#hystrix 1 0 0 0 1 0 0 0 Euonymus#alata 0 0 0 0 0 0 0 0 Euonymus#fortunei 96 12 0 0 70 12 0 0 Fraxinus#sp 280 125 174 126 133 85 74 51 Galium#aparine 18 4 86 26 18 4 81 26 Galium#circaezans 17 0 18 6 12 0 16 6 Galium#mollugo 0 0 26 4 0 0 19 4 Geranium#maculatum 0 3 0 0 0 3 0 0 Glechoma#hederacea 4 0 62 0 4 0 62 0 Impatiens#sp.# 24 73 48 88 24 45 48 51 Leersia#sp 46 32 142 84 23 21 55 74 Ligustrum#vulgare 80 556 40 30 80 374 40 30 Liriodendron#tulipfera 0 0 12 0 0 0 8 0 Lonicera#japonica 146 12 0 20 107 12 0 13 Lonicera#maackii 372 605 139 256 96 183 57 128 Ophioglossum#pusillum 2 6 4 9 2 6 4 9 Ostrya#virginiana 20 0 0 0 20 0 0 0 Parthenocissus#quinquefolia 249 153 377 114 167 59 175 56 Pilea#pumilla 29 0 77 204 20 0 25 114 Podophyllum#peltatum 0 0 200 60 0 0 89 60 Polygonum#sp. 0 0 0 16 0 0 0 12 Polyongatum#biflorum 104 6 22 14 62 6 16 14 Prunus#serotina 202 86 12 62 149 79 12 42 Quercus#sp.# 0 0 7 0 0 0 7 0 Ribes#cynosbati# 0 0 40 0 0 0 40 0 Rosa#multiflora 0 92 20 0 0 57 20 0 Sanguinaria#canadensis 60 0 0 0 40 0 0 0 Sanicula#canadensis 100 42 20 73 69 42 20 66 Sanicula#odorata 1247 844 270 183 1124 519 118 99 Setaria#lutescens 8 4 86 12 8 4 53 10 Smilax#sp. 46 24 24 30 39 24 17 23 Solanum#pumilum 50 40 48 12 31 40 30 12 Stellaria#media 0 0 4 0 0 0 4 0 Toxicodendron#radicans 0 12 4 0 0 12 4 0 Trillium#sp.# 0 0 39 2 0 0 39 2 Ulmus#rubra 86 50 28 64 53 50 15 44 Viola#sp. 34 0 140 0 32 0 78 0 Vitis#sp. 61 10 14 39 28 4 10 19 Unknown$Species$SUM2014F1 60 0 0 6 60 0 0 6 Unknown$Species$SUM2014F2g 44 10 21 14 31 10 17 14 Unknown$Species$SUM2014F3g 0 0 220 60 0 0 174 60 Unknown$Species$SUM2014F4 8 0 0 0 8 0 0 0 Unknown$Species$SUM2014F5 0 0 0 7 0 0 0 7 Unknown$Species$SUM2014F6 60 0 0 0 60 0 0 0 Unknown$Species$SUM2014F7 0 70 0 0 0 44 0 0 Unknown$Species$SUM2014F8g 40 0 0 8 40 0 0 8 *$g$after$the$unknown$species$number$refers$to$a$graminoidFlike$species
99! !
Appendix(Table(A3.!!Significance!of!treatment!effects!on!species!composition!of!forest!floor!plants!by!year!and!season.!!! Values!determined!using!permuatational!multivariate!analysis!of!variance.!!Significance!for!the!whole ! ! ! Sums!of! ! ! ! ! ! ! ! ! ! ! ! ! Square Mean!! Treatmen Sums!of! Perm Season! Treatment! Df! s! Squares! F.Model! R2! P! !! t! Df! Squares! F! s! P! Method:!Adonis! Method:!np.MANOVA! ! ! ! ! ! ! ! ! ! ! Spring!2011! Deer! ! 1! 0.2987! ! 0.29867! ! 1.8786! ! 0.06448! ! 0.014! ! ! Site! ! ! 4! 40700359! ! 2.87E+00! ! 999! ! 0.001! ! ! ! Honeysuckle! 1! 0.2089! 0.20895! 1.3143! 0.04511! 0.203! Deer! 5! 17757456! 1.60E+00! 999! 0.096! ! 0.5143 0.4441 0.02 ! 1 Site( 4( 2.0574( 4( 3.2352( 9( 9( Residuals! 0! 22193728! 2.22E+06! ! 1 0.4462 ! ! ! Residuals! 3! 2.0667! 0.15898! 2! ! 1 ! ! ! ! ! ! ! ! ! Total! 9! 4.6317! 1! Summer!2011! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! Deer! 1! 0.0918! 0.09184! 0.5544! 0.01779! 0.92! Site! 4! 21771492! 4.37E+00! 999! 0.001! ! Honeysuckle! 1! 0.1286! 0.12857! 0.7761! 0.02491! 0.709! ! Deer! 5! 6225145! 1.20E+00! 999! 0.174! ! ! 1 Site! 4! 2.788! 0.697! 4.2075! 0.54011! 0.922! Residuals! 0! 10375375! 1.04E+06! ! 1 0.4171 ! ! ! Residuals! 3! 2.1535! 0.16565! 9! ! 1 ! ! ! ! ! ! ! ! ! Total! 9! 5.1619! 1! Spring!2012! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! Deer! 1! 0.3923! 0.39227! 2.5697! 0.08034! 0.01! Site! 4! 45565797! 2.15E+00! 999! 0.045! ! ! 2648834 2.35E+0 0.02 Honeysuckle! 1! 0.1707! 0.17072! 1.1184! 0.03496! 0.34! Deer( 5( 0( 0( 999( 6( ! 0.5838 0.4782 0.01 ! 1 Site( 4( 2.3352( 1( 3.8244( 6( 7( Residuals! 0! 22543212! 2.25E+06! ! 1 0.4064 ! ! ! Residuals! 3! 1.9845! 0.15265! 3! ! ! ! ! ! ! ! ! ! ! 100! ! 1 Total! 9! 4.8827! 1! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! Appendix(Table(A3.!! Continued! Summer!2012! Deer! ! 1! 0.4589! ! 0.45891! ! 2.4293! ! 0.09287! ! 0.003! ! ! Site! ! ! 4! 15078317! ! 2.06E+00! ! 999! ! 0.024! ! ! ! Honeysuckle! 1! 0.2281! 0.22813! 1.2076! 0.04617! 0.26! Deer! 5! 9165579! 2.22E+00! 999! 0.114! ! 0.4495 0.3639 0.01 ! 1 Site( 4( 1.7983( 8( 2.3799( 5( 4( Residuals! 0! 8259512! 8.26E+05! ! 1 0.4970 ! ! ! Residuals! 3! 2.4558! 0.18891! 1! ! 1 ! ! ! ! ! ! ! ! ! Total! 9! 4.9412! 1! Spring!2013! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! Deer! 1! 0.4295! 0.42955! 2.0483! 0.07482! 0.014! Site! 4! 17922291! 2.11E+00! 999! 0.008! ! ! 1063187 2.26E+0 0.01 Honeysuckle! 1! 0.2449! 0.24491! 1.1679! 0.04266! 0.29! Deer( 5( 3( 0( 999( 5( ! 0.5850 0.4076 0.03 ! 1 Site( 4( 2.3401( 2( 2.7897( 3( 9( Residuals! 0! 9405151! 9.41E+05! ! 1 0.4748 ! ! ! Residuals! 3! 2.7262! 0.20971! 8! ! 1 ! ! ! ! ! ! ! ! ! Total! 9! 5.7408! 1! ! ! ! ! ! ! ! ! ! ! ! Summer!2013! Deer! ! 1! 0.4996! ! 0.4996! ! 2.0757! ! 0.0795! ! 0.007! ! ! Site! ! ! 4! 18293295! ! 1.49E+00! ! 999! ! 0.11! ! ! 0.4723 0.0751 0.02 ! Honeysuckle( 1( 0.4724( 8( 1.9626( 7( 1( Deer! 5! 15296618! 2.99E+00! 999! 0.121! ! 0.5457 0.00 ! 1 Site( 4( 2.183( 6( 2.2675( 0.3474( 2( Residuals! 0! 10224049! 1.02E+06! ! 1 0.4979 ! ! ! Residuals! 3! 3.129! 0.24069! 3! ! 1 ! ! ! ! ! ! ! ! ! Total! 9! 6.284! 1! ! ! ! ! ! ! ! ! ! ! ! Spring!2014! Deer! ! 1! 0.4361! ! 0.43614! ! 1.9927! ! 0.07794! ! 0.027! ! ! Site! ! ! 4! 17425208! ! 1.56E+00! ! 999! ! 0.121! ! ! ! 101! ! 1395661 2.10E+0 0.00 Honeysuckle! 1! 0.3081! 0.30811! 1.4077! 0.05506! 0.149! Deer( 5( 6( 0( 999( 3( ! 0.5016 0.3585 0.02 ! 1 Site( 4( 2.0065( 2( 2.2918( 5( 3( Residuals! 0! 13295856! 1.33E+06! ! 1 0.5084 ! ! ! Residuals! 3! 2.8453! 0.21887! 5! ! 1 ! ! ! ! ! ! ! ! ! Total! 9! 5.5961! 1! Summer!2014! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! Deer! 1! 0.6417! 0.64168! 3.3174! 0.11816! 0.001! Site! 4! 20382795! 1.45E+00! 999! 0.092! ! Honeysuckle! 1! 0.1864! 0.18644! 0.9639! 0.03433! 0.476! ! Deer! 5! 17552667! 2.82E+00! 999! 0.053! ! 0.5219 0.3844 0.00 ! 1 Site( 4( 2.0877( 3( 2.6983( 5( 1( Residuals! 0! 12459663! 1.25E+06! ! 1 0.4630 ! ! ! Residuals! 3! 2.5146! 0.19343! 6! ! 1 ! ! ! ! ! ! ! ! ! !! Total! 9! 5.4304! 1! !! !! !! ! ! ! ! ! ! ! 102! ! Appendix(Table(A4.!!Significance!of!treatment!types!on!cover!of!various!plant!function! types!in!species!and!years!with!significant!compositional!differences.!!Groups!that!were!not! analyzed!due!to!absence!in!≥!5!plots!are!not!shown.!!Significant!treatments!are!shown!in! bold.!! Season Year Functional/Type Treatment df den/df F P Spring 2012 Graminoid Deer 1 4 0.147335 0.7206 Honeysuckle 1 9 0.875376 0.3739 Spring/Perennial Deer 1 4 1.46701 0.2925 Honeysuckle 1 9 4.3889 0.0657 Summer/Perennial Deer 1 4 0.456813 0.5362 Honeysuckle 1 9 1.188162 0.304 Vine Deer 1 4 2.013942 0.2289 Honeysuckle 1 9 0.188242 0.6746 Shrub Deer 1 4 0.010647 0.9228 Honeysuckle 1 9 9.034911 0.0148 Tree Deer 1 4 1.18385 0.3377 Honeysuckle 1 9 1.392539 0.2682 Annual Deer 1 4 1.3473249 0.3103 Honeysuckle 1 9 0.2603649 0.6221 Spring 2013 Graminoid Deer 1 4 2.128206 0.2184 Honeysuckle 1 9 4.537202 0.062 Spring/Perennial Deer 1 4 3.11823 0.1522 Honeysuckle 1 9 6.8994 0.0275 Summer/Perennial Deer 1 4 0.548472 0.5001 Honeysuckle 1 9 0.490821 0.5013 Vine Deer 1 4 0.057635 0.8221 Honeysuckle 1 9 0.765421 0.4044 Shrub Deer 1 4 0.001966 0.9668 Honeysuckle 1 9 4.610295 0.0603 Tree Deer 1 4 4.974043 0.0896 Honeysuckle 1 8 4.756626 0.0608 Deer:Honeysuckle 1 8 14.216586 0.0055 Annual Deer 1 4 0.9429 0.3865 Honeysuckle 1 9 7.188848 0.0252 ! ! Spring 2014 Graminoid Deer 103! 1 4 2.511832 0.1882 Honeysuckle 1 9 10.457304 0.0103 Spring/Perennial Deer 1 4 1.934583 0.2366 Honeysuckle 1 9 4.644981 0.0595 Summer/Perennial Deer 1 4 0.8089714 0.4193 Honeysuckle 1 9 0.7200323 0.4181 Vine Deer 1 4 0.042014 0.8476 Honeysuckle 1 9 4.336452 0.067 Shrub Deer 1 4 0.377925 0.572 Honeysuckle 1 9 4.494003 0.063 Tree Deer 1 4 6.98087 0.0575 Honeysuckle 1 9 5.569606 0.0426 Biennial Deer 1 4 2.67886 0.177 Honeysuckle 1 9 1.340441 0.2768 Summer 2013 Graminoid Deer 1 4 0.562583 0.4949 Honeysuckle 1 9 2.974144 0.1187 Summer/Perennial Deer 1 4 0.5147536 0.5128 Honeysuckle 1 9 0.4847632 0.5039 Vine Deer 1 4 0.004936 0.9474 Honeysuckle 1 9 16.79624 0.0027 Shrub Deer 1 4 0.241795 0.6487 Honeysuckle 1 9 3.622034 0.0894 Tree Deer 1 4 2.072527 0.2234 Honeysuckle 1 8 0.793158 0.3991 Deer:Honeysuckle 1 8 5.967552 0.0404 Season Year Functional/Type Treatment df den/df F P Spring 2012 Graminoid Deer 1 4 0.147335 0.7206 Honeysuckle 1 9 0.875376 0.3739 Spring/Perennial Deer 1 4 1.46701 0.2925 Honeysuckle 1 9 4.3889 0.0657 Summer/Perennial Deer 1 4 0.456813 0.5362 Honeysuckle 1 9 1.188162 0.304 Vine Deer 1 4 2.013942 0.2289 Honeysuckle 1 9 0.188242 0.6746 Shrub Deer 1 4 0.010647 0.9228 Honeysuckle 1 9 9.034911 0.0148 Tree Deer 1 4 1.18385 0.3377 Honeysuckle 1 9 1.392539 0.2682 Annual Deer 1 4 1.3473249 0.3103 Honeysuckle 1 9 0.2603649 0.6221 Spring 2013 Graminoid Deer 1 4 2.128206 0.2184 Honeysuckle 1 9 4.537202 0.062 Spring/Perennial Deer 1 4 3.11823 0.1522 Honeysuckle 1 9 6.8994 0.0275 Summer/Perennial Deer 1 4 0.548472 0.5001 Honeysuckle 1 9 0.490821 0.5013 Vine Deer 1 4 0.057635 0.8221 Honeysuckle 1 9 0.765421 0.4044 Shrub Deer 1 4 0.001966 0.9668 Honeysuckle 1 9 4.610295 0.0603 ! Tree Deer 1 4 4.974043 0.0896 Honeysuckle 1 8 4.756626 0.0608 Appendix(Table(A4!Continued.Deer:Honeysuckle! 1 8 14.216586 0.0055 Annual den/df Season Year Functional/Type TreatmentDeer df1 4 0.9429F 0.3865P ! Spring 2012 Honeysuckle 1 9 7.188848 0.0252 Spring! ! 2014 Graminoid! ! Graminoid Deer 1 4 0.147335 0.7206 HoneysuckleDeer 1 94 0.8753762.511832 0.37390.1882 Spring/Perennial Honeysuckle 1 9 10.457304 0.0103 Spring/Perennial Deer 1 4 1.46701 0.2925 HoneysuckleDeer 1 94 1.9345834.3889 0.06570.2366 Summer/Perennial Honeysuckle 1 9 4.644981 0.0595 Summer/Perennial Deer 1 4 0.456813 0.5362 HoneysuckleDeer 1 94 0.80897141.188162 0.41930.304 Vine Honeysuckle 1 9 0.7200323 0.4181 Vine Deer 1 4 2.013942 0.2289 HoneysuckleDeer 1 94 0.1882420.042014 0.67460.8476 Shrub Honeysuckle 1 9 4.336452 0.067 Shrub Deer 1 4 0.010647 0.9228 HoneysuckleDeer 1 94 9.0349110.377925 0.01480.572 Tree Honeysuckle 1 9 4.494003 0.063 Tree Deer 1 4 1.18385 0.3377 HoneysuckleDeer 1 94 1.3925396.98087 0.26820.0575 Annual Honeysuckle 1 9 5.569606 0.0426 Deer 1 4 1.3473249 0.3103 Biennial Honeysuckle 1 9 0.2603649 0.6221 Spring 2013 Deer 1 4 2.67886 0.177 Graminoid Honeysuckle 1 9 1.340441 0.2768 Summer 2013 Deer 1 4 2.128206 0.2184 Graminoid Honeysuckle 1 9 4.537202 0.062 Spring/Perennial Deer 1 4 0.562583 0.4949 DeerHoneysuckle 1 49 2.9741443.11823 0.15220.1187 Summer/Perennial Honeysuckle 1 9 6.8994 0.0275 Summer/Perennial Deer 1 4 0.5147536 0.5128 DeerHoneysuckle 1 49 0.48476320.548472 0.50010.5039 Vine Honeysuckle 1 9 0.490821 0.5013 Vine Deer 1 4 0.004936 0.9474 DeerHoneysuckle 1 49 0.05763516.79624 0.82210.0027 Shrub Honeysuckle 1 9 0.765421 0.4044 Shrub Deer 1 4 0.241795 0.6487 DeerHoneysuckle 1 49 0.0019663.622034 0.96680.0894 Tree Honeysuckle 1 9 4.610295 0.0603 Tree Deer 1 4 2.072527 0.2234 DeerHoneysuckle 1 48 4.9740430.793158 0.08960.3991 HoneysuckleDeer:Honeysuckle 1 8 4.7566265.967552 0.06080.0404 Deer:Honeysuckle 1 8 14.216586 0.0055 ! Annual ! ! Deer 1 4 0.9429 0.3865 Honeysuckle 1 9 7.188848 0.0252 Spring 2014 Graminoid Deer 1 4 2.511832 0.1882 Honeysuckle 1 9 10.457304 0.0103 Spring/Perennial Deer 1 4 1.934583 0.2366 Honeysuckle 104! 1 9 4.644981 0.0595 Summer/Perennial Deer 1 4 0.8089714 0.4193 Honeysuckle 1 9 0.7200323 0.4181 Vine Deer 1 4 0.042014 0.8476 Honeysuckle 1 9 4.336452 0.067 Shrub Deer 1 4 0.377925 0.572 Honeysuckle 1 9 4.494003 0.063 Tree Deer 1 4 6.98087 0.0575 Honeysuckle 1 9 5.569606 0.0426 Biennial Deer 1 4 2.67886 0.177 Honeysuckle 1 9 1.340441 0.2768 Summer 2013 Graminoid Deer 1 4 0.562583 0.4949 Honeysuckle 1 9 2.974144 0.1187 Summer/Perennial Deer 1 4 0.5147536 0.5128 Honeysuckle 1 9 0.4847632 0.5039 Vine Deer 1 4 0.004936 0.9474 Honeysuckle 1 9 16.79624 0.0027 Shrub Deer 1 4 0.241795 0.6487 Honeysuckle 1 9 3.622034 0.0894 Tree Deer 1 4 2.072527 0.2234 Honeysuckle 1 8 0.793158 0.3991 Deer:Honeysuckle 1 8 5.967552 0.0404 ! ! Appendix(Table(A5.(SplitBplot!ANOVA!statistics!for!the!significance!of!treatment!effects!on! cover!of!nonBLonicera!maackii!invasive!plants!by!year!and!season.!!!For!each!model,!the! Deer*Honeysuckle!interaction!term!was!not!significant,!and!was!dropped!from!the!model.! ! Season Year Treatment numDF denDF F0value p0value Spring 2011 Deer 1 13 0.07252 0.7919 Honeysuckle 1 13 0.91035 0.3574 Spring 2012 Deer 1 4 0.40807 0.5577 Honeysuckle 1 9 0.23321 0.6407 Spring 2013 Deer 1 4 0.02203 0.8892 Honeysuckle 1 9 0.28746 0.6048 Spring 2014 Deer 1 4 0.01269 0.9157 Honeysuckle 1 9 0.23827 0.6371 Summer 2011 Deer 1 13 1.22335 0.2888 Honeysuckle 1 13 0.64342 0.4369 Summer 2012 Deer 1 13 0.90346 0.3592 Honeysuckle 1 13 0.00118 0.9731 Summer 2013 Deer 1 13 3.24293 0.0950 Honeysuckle 1 13 1.44429 0.2509 Summer 2014 Deer 1 13 0.81037 0.3844 Honeysuckle 1 13 0.05729 0.8146 ! ! 105! ! ! Appendix!Table(A6.(ANOVA!Table!for!cover!of!Sanicula!odorata!and!Alliaria!petiolata!in! summer!of!2011.( ! Species! Effect! numDF! denDF& F&& P& Sanicula&odorata&1&Summer&2011& Deer! 1! ! 4! !0.324187! ! 0.5996! ! Honeysuckle! 1! 9! 0.833014! 0.3852! Alliaria&petiolata&! 1&Summer&2011& Deer! 1! ! 4! !0.068402! ! 0.8066! ! ! ! ! 106! ! Appendix!Figure(A1.!!Schematic!of!sampling!locations!within!each!deer!exclosure!or!deer! access!half!plot.!!!L.m.!is!an!abbreviation!for!Lonicera&maackii.!!Each!transect!(blue!line)!had! a!sampling!location!(small!blue!rectangles)!every!2m.!!! ! ! 107! ! ! Appendix(Figure(A10.!!Spring!plant!species!richness!by!time.!!Graph!shows!mean!plant! species!richness,!+/B!SE,!across!4!years.!Areas!with!deer!excluded!are!represented!with!a! square,!while!areas!with!deer!access!are!represented!with!a!circle.!!Areas!with!Lonicera! maackii!removal!are!noted!with!an!open!symbol,!areas!with!L.!maackii!left!intact!are!noted! with!a!closed!symbol.! ! ! 108! ! ! ! Appendix(Figure(A3.!!NMDS!ordinations!of!species!composition!data!by!season!and!year.!! Site!is!represented!by!color.!!Open!symbols!represent!areas!of!deer!exclosure,!while!closed! symbols!represent!areas!of!deer!access.!!Squares!represent!areas!of!Lonicera&maackii&(L.m.)! removal,!while!triangles!represent!areas!of!L.&maackii&presence.! ! 109! ! Summer Species Richness 2011 - 2014 17 Deer EX AC 16 15 14 Species Richness Species 13 12 11 PR RE Honeysuckle Treatment ( ! Appendix(Figure(A4.((Interaction!plot!of!summer!plant!species!richness!interaction!plot!of! forest!floor!plant!species!richness,!based!on!summer!samples,!showing!mean!+B!SE!for!each! treatment.!!Open!circles!represent!areas!of!deer!access;!filled!circles!represent!areas!with! deer&exclosures.!!Lonicera&maackii!treatment!is!shown!on!the!xBaxis,!with!areas!with!L.& maackii&presence!marked!as!PR,!and!areas!that!have!L.&maackii!removed!marked!as!RE.!! ! ! 110! ! Additional(Tables(and(Figures(from(Chapter(3( ! Appendix(Table(A1.!!Tree!seedling!planting,!removal!and!sampling!dates.!!Check!1!was!only!of! survival,!not!height!or!browse!status.! ! Event! Date!or!Date!Range! Seedling!Planting! 5/31/14!B!6/6/14! Check!1! 6/19/14! Check!2! 7/3/14! Check!3! 8/7/14! Check!4! 9/7/14!B!9/9/14! Check!5! 10/5/14!B!10/8/14! Check!6! 12/17/14!B!12/18/14! Check!7! 4/1/15! Check!8! 5/5/15!B!5/7/15! Check!9! 6/11/15!B!6/14/15! Seedling!Removal! 6/23/15!B!6/25/15! ! 111! ! Appendix(Table(2A.!Seedling!mortality!by!treatment!type!and!species.!X2"values!were!calculated!from!weighted!observed!survival!and" weighted!Kaplan=Meier!estimates!of!predicted!survival,!and!differences!between!treatments!were!calculated!using!the!Peto!and!Peto! modification!of!the!Gehan=Wilcoxon!test(Pyke!and!Thompson!1986,!Rich!et!al.!2010).!!O!is!the!weighted!observed!survival,!E!is!the!expected! survival!and!V!is!variance.!Significant!effects!are!bolded.!!! ! Species! Treatment! Chisq! df" P! !! Location! N! Observed! Expected! (O=E)^2/E! (O=E)^2/V! ACSA! Deer! ( ! 4.2!( 1(! 0.0407!( Access! ! 20! ! 10.23! ! 6.87! ! 1.64! ! 4.19! ! ! ! Exclosure! 19! 4.31! 7.67! 1.47! 4.19! ! Honeysuckle! ! ! 0.7! 1! 0.409! Away! 19! 8.36! 7! 0.264! 0.683! ! ! Under! 20! 6.18! 7.54! 0.245! 0.683! ! Deer:Honeysuckle! ( ! 12.7!( 3(! 0.00545!( AccessAway! 10! 7.59! 3.08! 6.6188! 11.0704! ! ! AccessUnder! 10! 2.641! 3.79! 0.3508! 0.6381! ! ! ! ! ! ! ExclosureAway! 9! 0.769! 3.92! 2.5354! 4.7398! ExclosureUnder! 10! 3.538! 3.74! 0.0112! 0.0204! CAOV! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! Deer! 2.2! 1! 0.137! Access! 20! 10.05! 7.65! 0.756! 2.22! ! ! Exclosure! 17! 5.86! 8.27! 0.7! 2.22! ! ! ! ! ! ! Honeysuckle! 0.7! 1! 0.405! Away! 18! 6.35! 7.7! 0.237! 0.695! ! ! Under! 19! 9.57! 8.22! 0.222! 0.695! ! ! ! ! ! ! Deer:Honeysuckle! 3.2! 3! 0.365! AccessAway! 10! 4.68! 3.81! 0.19628! 0.3862! ! ! AccessUnder! 10! 5.38! 3.84! 0.61839! 1.2174! ! ! ! ! ! ! ExclosureAway! 8! 1.68! 3.89! 1.26201! 2.5649! ! ! ! ! ! ! ExclosureUnder! 9! 4.19! 4.38! 0.00817! 0.0173! ! ! ! ! ! ! QUAL! Deer! ! ! 0! 1! 0.971! Access! ! 20! ! 1!! 0.975! ! 0.000641! ! 0.00128! ! ! ! Exclosure! 20! 0.975! 1! 0.000625! 0.00128! ! Honeysuckle! ! ! 2.1! 1! 0.152! Away! 20! 0! 1! 1! 2.05! ! ! Under! 20! 1.98! 0.975! 1.03! 2.05! ! Deer:Honeysuckle! ! ! 2.1! 3! 0.561! AccessAway! 10! 0! 0.5! 0.5! 0.678! ! ! AccessUnder! 10! 1! 0.475! 0.58! 0.774! ! ! ! ! ! ! ExclosureAway! 10! 0! 0.5! 0.5! 0.678! ! ! ! ! ! ! !! !! !! !! !! !! ExclosureUnder! 10! 0.975! 0.5! 0.451! 0.612! 112! ! Appendix(Table(3A.!!Significance!of!treatment!effects!on!species!browse!status!by!species!for!seedlings!planted!in! deer!access!plots.!!Table!shows!results!from!a!split;plot,!generalized!linear!mixed;effect!model,!modified!for!binary! data!with!a!binomial!distribution.!Significance!of!effects!were!determined!by!maximum!likelihood!(Laplace! Approximation).!! ! Species! Treatment! Estimate! Std.!Error! z!value! Pr(>|z|)! ACSA! Honeysuckle! ! ! 1.40! ! 1.03! ! 1.359! ! 0.174! CAOV! ! !! Honeysuckle! ! ! ;0.4055! ! 0.9037! ! ;0.449! ! 0.654! ! ! 113! ! Appendix(Table(4A.!!Significance!of!treatment!effects!on!root!mass,!shoot!mass!and!root:shoot!ratio!of!tree! seedlings.!Values!determined!using!split;plot!analysis!of!variance.!!Bolded!values!are!significant!at!α!=!0.05.!! ! ! ! Response Species Treatment df den0df F P Root0Mass ACSA Site 1 16 5.7486 0.0291 Deer 1 16 2.2045 0.1570 Honeysuckle 1 8 1.0127 0.3437 CAOV Site 1 15 1.1387 0.3028 Deer 1 15 3.0003 0.1038 Honeysuckle 1 7 3.2736 0.1133 QUAL Site 1 17 0.5638 0.4630 Deer 1 17 0.1000 0.7556 Honeysuckle 1 17 0.0453 0.8340 Shoot0Mass ACSA Site 1 16 4.8638 0.0424 Deer 1 16 2.1826 0.1590 Honeysuckle 1 8 0.0584 0.8151 CAOV Site 1 15 4.2326 0.0575 Deer 1 15 2.3264 0.1480 Honeysuckle 1 7 0.0036 0.9537 QUAL Site 1 17 1.8758 0.1886 Deer 1 17 1.0574 0.3182 Honeysuckle 1 17 1.3003 0.2700 Root:Shoot0Ratio ACSA Site 1 16 1.4318 0.2489 Deer 1 16 0.0634 0.8044 Honeysuckle 1 8 0.7902 0.4000 CAOV Site 1 15 2.5214 0.1332 Deer 1 15 0.0798 0.7814 Honeysuckle 1 7 3.8168 0.0917 QUAL Site 1 17 0.0413 0.8413 Deer 1 17 4.5570 0.0476 Honeysuckle 1 17 3.7448 0.0698 114! ! CAOV Survival - Kaplan-Meier Curve 1.0 0.8 + 0.6 + + + 0.4 Overall Survival Proportion Survival Overall AccessAway AccessUnder ExclosureAway 0.2 ExclosureUnder 0.0 0 10 20 30 40 50 Time (Weeks) ! ( Appendix(Figure(1A.!Survival!Curves!for!Carya%ovata!(CAOV)%survival!over!the!course!of!the!50!week!experiment.! ! ! 115! ! QUAL Survival - Kaplan-Meier Curve + 1.0 + 0.8 0.6 0.4 Overall Survival Proportion Survival Overall AccessAway AccessUnder ExclosureAway 0.2 ExclosureUnder 0.0 0 10 20 30 40 50 Time (Weeks) ! ! Appendix(Figure(2A.!!Survival!Curves!for!Quercus%alba%(QUAL)%survival!over!the!course!of!the!50!week! experiment.! ! ! 116! ! ACSA CAOV QUAL 100 75 Deer 50 ACC Browsed (Percent) Browsed 25 0 Away Under Away Under Away Under Honeysuckle ( Appendix(Figure(3A.!!Percent!browse!(+/;!SE)!of!Acer%saccharum%(ACSA),!Carya%ovata!(CAOV),!and!Quercus%alba! (QUAL)!seedlings!planted!under!or!away!from!the!canopy!of!L.%maackii!and!with!(ACC)!deer!access.!!! ! ! 117! ! ACSA CAOV QUAL 7.5 Deer 5.0 Exclosure Access Root Mass (g) Root 2.5 0.0 Away Under Away Under Away Under Honeysuckle ! ! Appendix(Figure(4A.!!Root!mass!of!Acer%saccharum%(ACSA),!Carya%ovata!(CAOV),!and!Quercus%alba!(QUAL)! seedlings!planted!under!or!away!from!the!canopy!of!L.%maackii!and!with!(ACC)!or!without!(EXC)!deer!access.!!! ! ! 118! ! Additional(Table(from(Chapter(4( ( Appendix(Table(A1.!!Paired!t;tests!for!each!height!range!and!individual!height!between!deer!exclosures! and!access!plots.!!Significant!differences!are!bolded.!! ! Category! Height! df! t! p! Range! 0!;!0.5!m!! ! 4!! ;0.885! ! 0.4263! ! ! 0.5(>(1(m(( 4( >2.806( 0.0485( ! 1(>(1.5(m(( 4( >5.729( 0.0046( !Individual!Heights! 0.5!m!! 4!! ;2.298! 0.0832! ! ! 1(m(( 4( >4.882( 0.0081( ! 1.5!m!! 4! ;0.233! 0.8276! !! 2!m!! 4! 0.739! 0.5012! ( ! ! ! ! ! 119!