Interactions between a Gall Making Fly, Dasineura ...
Transcript of Interactions between a Gall Making Fly, Dasineura ...
University of Massachusetts Amherst University of Massachusetts Amherst
ScholarWorks@UMass Amherst ScholarWorks@UMass Amherst
Open Access Dissertations
2-2013
Interactions between a Gall Making Fly, Dasineura Oxycoccana Interactions between a Gall Making Fly, Dasineura Oxycoccana
(Diptera: Cecidomyiidae), And Its Host Plant, Cultivated Cranberry (Diptera: Cecidomyiidae), And Its Host Plant, Cultivated Cranberry
(Vaccinium Macrocarpon) (Vaccinium Macrocarpon)
Sunil Tewari University of Massachusetts Amherst
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Part of the Entomology Commons
Recommended Citation Recommended Citation Tewari, Sunil, "Interactions between a Gall Making Fly, Dasineura Oxycoccana (Diptera: Cecidomyiidae), And Its Host Plant, Cultivated Cranberry (Vaccinium Macrocarpon)" (2013). Open Access Dissertations. 710. https://doi.org/10.7275/etrn-a140 https://scholarworks.umass.edu/open_access_dissertations/710
This Open Access Dissertation is brought to you for free and open access by ScholarWorks@UMass Amherst. It has been accepted for inclusion in Open Access Dissertations by an authorized administrator of ScholarWorks@UMass Amherst. For more information, please contact [email protected].
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!! ,)D&!!ACKNOWLEDGMENTS ................................................................................................... v
ABSTRACT ....................................................................................................................... vi
LIST OF TABLES ............................................................................................................. xi
LIST OF FIGURES ........................................................................................................... xii
CHAPTER
1. INJURY TO APICAL MERISTEM OF CRANBERRY BY DASINEURA
OXY COCCANA (DIPTERA: CECIDOMYIIDAE) REDUCES
PRODUCTION OF FLORAL-UNITS IN THE NEXT GROWING
SEASON ................................................................................................................. 1
!Introduction ................................................................................................................... 1
Materials and Methods .................................................................................................. 6
Upright-level Effect: Tagging Study ....................................................................... 6
Site-level Effect: Sampling Study ........................................................................... 8
Plot-level Effect: Suppression Study ..................................................................... 10
Data analysis ......................................................................................................... 11
!Results ......................................................................................................................... 14
!Upright-level Effect: Tagging Study (Massachusetts) .......................................... 14
Upright-level Effect: Tagging Study (Maine) ....................................................... 18
Site-level Effect: Sampling Study (Massachusetts) .............................................. 19
Site-level Effect: Sampling Study (Maine) ........................................................... 23
Plot-Level Effect: Suppression Study ................................................................... 25
!Discussion ................................................................................................................... 26
!2. IMPACT OF EARLY SEASON APICAL MERISTEM INJURY BY
GALL-INDUCING, DASINEURA OXY COCCANA (DIPTERA:
CECIDOMYIIDAE), ON REPRODUCTIVE AND VEGETATIVE
GROWTH OF CRANBERRY .............................................................................. 36
!Introduction ................................................................................................................. 36
Materials and Methods ................................................................................................ 40
!Upright-level field study (Year 1: 2010) ............................................................... 40
!
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Upright-level field study (Year 2: 2011) ............................................................... 42
Greenhouse Study ................................................................................................. 42
Data analysis ......................................................................................................... 44
!Results ......................................................................................................................... 46
!Upright-level field study (Year 1: 2010): Impact of early season apical
meristem injury on number of flowers, fruit set, and fruit weight .................. 46
Production of side-shoots in response to early seson tipworm and simulted
apical meristem injury ..................................................................................... 48
Upright-level field study (Year 2: 2011): Impact of apical meristem injury
on vegetative growth and flowering in the next growing season .................... 49
Greenhouse study .................................................................................................. 53
!Discussion ................................................................................................................... 53
!3. PHYSIOLOGICAL INTEGRATION PLAYS KEY ROLE IN
CRANBERRY TOLERANCE OF DAMAGE BY TIPWORM LARVAE,
DASINEURA OXY COCCANA (DIPTERA: CECIDOMYIIDAE) .................... 61
!Introduction ................................................................................................................. 61
Materials and Methods ................................................................................................ 66
!Determining the role of current season growth in tolerance of apical
meristem injury (Field Study) ......................................................................... 66
Effect of girdling on growth and movement of assimilates in intact
cranberry uprights (supplement of field study) ............................................... 67
Data analysis ......................................................................................................... 69
!Results ......................................................................................................................... 71
!Determining the role of physiological integration in the tolerance of
tipworm feeding injury .................................................................................... 71
Effect of girdling on growth and movement of assimilates in intact
cranberry uprights ........................................................................................... 74
!Discussion ................................................................................................................... 75
4. DEVELOPING FRUITS INHIBIT THE REGROWTH OF CRANBERRY
SHOOTS AFTER APICAL MERISTEM INJURY BY LARVAE OF
DASINEURA OXY COCCANA (DIPTERA: CECIDOMYIIDAE) .................... 80
!Introduction ................................................................................................................. 80
Materials and Methods ................................................................................................ 83
!
!
]!
Comparing flowering and vegetative uprights for the production of side-
shoots after tipworm injury (Field Study 1) .................................................... 84
Determining impact of flower bud removal on the production of side-
shoots in tipworm injured flowering uprights (Field Study 2) ........................ 84
Determining impact of flower bud removal on the production of side-
shoots in tipworm injured flowering uprights (Greenhouse Study) ................ 85
Data analysis ......................................................................................................... 86
Results ......................................................................................................................... 87
!Comparing flowering and vegetative uprights for the production of side-
shoots after tipworm injury (Field Study 1) .................................................... 87
Determining impact of flower bud removal on the production of side-
shoots in tipworm injured flowering uprights (Field Study 2) ........................ 88
Determining impact of flower bud removal on the production of side-
shoots in tipworm injured flowering uprights (Greenhouse Study) ................ 89
!Discussion ................................................................................................................... 90
!5. THE EFFECT OF INJURY TIME AND CULTIVAR ON THE
TOLERANCE OF SIMULATED APICAL MERISTEM HERBIVORY
IN CRANBERRY ................................................................................................. 96
!Introduction ................................................................................................................. 96
Materials and Methods ................................................................................................ 99
!Data analysis ....................................................................................................... 101
!Results ....................................................................................................................... 102
!Vegetative regrowth - side-shoots (2010) ........................................................... 102
Fruit set (2010) .................................................................................................... 103
Vegetative growth of uprights (2011) ................................................................. 104
Flowering (2011) ................................................................................................. 105
!Discussion ................................................................................................................. 107
!BIBLIOGRAPHY ........................................................................................................... 110
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MA3
Proportion of uprights with flowers
0.20 0.22 0.24 0.26 0.28 0.30 0.32 0.34 0.36
Pro
port
ion o
f u
pri
ghts
wit
h f
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0.18
0.20
0.22
0.24
0.26
0.28
0.30
0.32
0.34
0.36
MA7
Proportion of uprights with flowers
0.25 0.30 0.35 0.40 0.45 0.50 0.55
Pro
port
ion o
f u
pri
ghts
wit
h f
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0.15
0.20
0.25
0.30
0.35
0.40
0.45
0.50
0.55
MA5
Proportion of uprights with flowers
0.40 0.45 0.50 0.55 0.60 0.65 0.70 0.75 0.80
Pro
port
ion o
f u
pri
ghts
wit
h f
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0.15
0.20
0.25
0.30
0.35
0.40
0.45
0.50
0.55 MA6
Proportion of uprights with flowers
0.26 0.28 0.30 0.32 0.34 0.36 0.38 0.40 0.42 0.44
Pro
port
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f u
pri
ghts
wit
h f
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0.10
0.15
0.20
0.25
0.30
0.35
0.40
MA8
Proportion of uprights with flowers
0.25 0.30 0.35 0.40 0.45 0.50 0.55 0.60 0.65
Pro
port
ion o
f upri
ghts
wit
h f
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0.0
0.1
0.2
0.3
0.4
0.5
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ME1
Proportion of uprights with flowers
0.08 0.10 0.12 0.14 0.16 0.18 0.20
Pro
port
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of
up
rig
hts
wit
h f
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0.04
0.06
0.08
0.10
0.12
0.14
0.16
0.18
0.20
0.22
0.24
ME2
Proportion of uprights with flowers
0.04 0.06 0.08 0.10 0.12 0.14 0.16 0.18 0.20 0.22 0.24
Pro
po
rtio
n o
f u
pri
gh
ts w
ith
fru
it
0.00
0.05
0.10
0.15
0.20
0.25
ME5
Proportion of uprights with flowers
0.05 0.10 0.15 0.20 0.25 0.30 0.35 0.40 0.45
Pro
po
rtio
n o
f u
pri
gh
ts w
ith
fru
it
0.0
0.1
0.2
0.3
0.4
0.5
ME6
Proportion of uprights with flowers
0.18 0.20 0.22 0.24 0.26 0.28 0.30 0.32 0.34
Pro
port
ion o
f u
pri
ghts
wit
h f
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0.14
0.16
0.18
0.20
0.22
0.24
0.26
0.28
0.30
0.32
0.34
0.36
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7&!%9DD&%(!(;)(!>*+D[(&':!%(9-$&%!$+!O;$=;!($BO*':!-&+%$($&%!)'&!&]B&'$:&+()>>/!
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%;&-!:*'&!>$D;(!*+!(;&!+)(9'&!*?!(*>&')+=&!&];$.$(&-!./!?>*O&'$+D!9B'$D;(%!*?!
=')+.&''/R!!
!
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45!6#$%&7*'$'!5(/2!!5&$8J*
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$%!:)-&!B*%%$.>&!./!(;&!:&():&'$=!)'=;$(&=(9'&!*?!B>)+(%!ZA)'C9$%!L__e\R!
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%9??$=$&+(!'&%*9'=&%!(*!(;&!-*':)+(!.9-%!ZU$>.'*9D;!)+-!8$=;)'-%!L__M\R!!
5;&!=9>($@)(&-!=')+.&''/S!."--$%$'/)/"-(*-"(0*%!"$(*+S!$%!)!>*O[D'*O$+D!
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!
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m9&.&=!Z"D'$=9>(9'&!)+-!"D'$[G**-!F)+)-)!JKK_\R!5$BO*':!$%!)!D)>>!:)V$+D!?>/!(;)(!
*@&'O$+(&'%!)%!>)(&!$+%()'!>)'@)&!Z<)D+&!L_^_\!)+-!)-9>(%!&:&'D&!)'*9+-!:$-[A)/!
$+!A)%%)=;9%&((%R!5$BO*':!?&:)>&%!>)/!*+&!*'!:*'&!&DD%!*+!(;&!)=($@&>/!D'*O$+D!
=')+.&''/!9B'$D;(%!)+-!(;&!>)'@)&!V$>>!(;&!($B!./!B$&'=$+D!(;'*9D;!(;&!:&'$%(&:)($=!
($%%9&!ZF**V!&(!)>R!JKLLh!G$D%R!L^!)+-!L_\R!5;&!>$?&!=/=>&!?'*:!&DD%!(*!)-9>(%!()V&%!LK[
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"?(&'!($BO*':!?&&-$+D!$+W9'/S!=')+.&''/!9B'$D;(%!:)/!'&%9:&!@&D&()($@&!
D'*O(;!./!(;&!)=($@)($*+!*?!>)(&')>!:&'$%(&:%!)+-!?*':!)!%$-&[%;**(!ZG$D%R!L^!)+-!
L_\R!H*O&@&'S!(;&!:)W*'$(/!*?!($BO*':[$+W9'&-!?>*O&'$+D!9B'$D;(%!-*!+*(!B'*-9=&!)!
%$-&[%;**(!.&?*'&!(;&!*+%&(!*?!-*':)+=/!)+-!D'*O(;!Z@&D&()($@&!)+-!'&B'*-9=($@&\!
$+!(;&!?*>>*O$+D!/&)'!:)/!.&!)-@&'%&>/!)??&=(&-!Z0!5&O)'$!9+B9.>$%;&-!-)()\R!3(!$%!
/&(!(*!.&!-&(&':$+&-!$?!>)=V!*?!)=($@&!:&'$%(&:%!Z7;$(;):!&(!)>R!L__L\!B'&@&+(%!
'&D'*O(;!)?(&'!($BO*':!?&&-$+D!$+W9'/S!*'!O;&(;&'!B'&%&+=&!*?!-&@&>*B$+D!?'9$(%!
$+;$.$(%!(;&!B'*-9=($*+!*?!%$-&[%;**(%!(;'*9D;!%$+V[%$+V!=*:B&($($*+!ZH*!L_^^\!$+!
?>*O&'$+D!=')+.&''/!9B'$D;(%R!E9'!%(9-/!)--'&%%&%!(;&!?*>>*O$+D!C9&%($*+%P!L\!3%!
(;&'&!)!-$??&'&+=&!.&(O&&+!($BO*':[$+W9'&-!@&D&()($@&!)+-!?>*O&'$+D!9B'$D;(%!$+!
(;&!B'*-9=($*+!*?!%$-&[%;**(%~!)+-!J\!#*&%!(;&!B'&%&+=&!*?!-&@&>*B$+D!?'9$(%!
%9BB'&%%!'&D'*O(;!)?(&'!($BO*':!?&&-$+D!$+W9'/~!
!
^J!
**
1GHEBI*<ZJ*;IHIANAGUI*FBN@OIBBP*EQBGHRA*MGAR*AGQMCBS*KIIDG@H*G@VEBP*N@D*N*
TGDI>TRCCA*QBCDEFID*KBCS*LNAIBNL*N\GLLNBP*OEDJ***
!
!
^M!
!!1GHEBI*<[J*1LCMIBG@H*FBN@OIBBP*EQBGHRA*MGAR*AGQMCBS*KIIDG@H*G@VEBP3*TGDI>
TRCCA*QBCDEFID*G@*BITQC@TI*AC*ARI*G@VEBP3*N@D*DIUILCQG@H*KBEGAT*G@*ONTGQIANL*
QCTGAGC@TJ**
*
/NAIBGNLT*N@D*/IARCDT*
General methods (Field studies 1 and 2). The field work was carried out at a
commercial cranberry farm with established beds of Howes and Stevens (41°56'59.31" N
70°47'15.98" W), planted in 1985 and 1995, respectively. For each cultivar, two 0.2 ha
plots were established and four 10 m equidistant transects were marked (in each plot).
The transects were laid down perpendicular to the direction in which the majority of
runners was oriented to eliminate measuring more than one upright on a single runner.
!
^g!
'CSQNBG@H*KLCMIBG@H*N@D*UIHIANAGUI*EQBGHRAT*KCB*ARI*QBCDEFAGC@*CK*TGDI>
TRCCAT*NKAIB*AGQMCBS*G@VEBP*41GILD*)AEDP*<8*
!E+&!B>*(!$+!&)=;!*?!(;&!=9>($@)'!.&-%!O)%!9%&-!ZH*O&%!)+-!0(&@&+%\R!G*'!&)=;!
(')+%&=(!$+!.*(;!(;&!=9>($@)'!B>*(%S!Ld!?>*O&'$+D!9B'$D;(%!)+-!)+!&C9)>!+9:.&'!*?!
@&D&()($@&!9B'$D;(%S!)>>!O$(;!@$%$.>&!($BO*':!?&&-$+D!$+W9'/!%$D+%!Z=9BB$+D!*?!
(&':$+)>!>&)@&%!O$(;!-&)-!)B$=)>!:&'$%(&:\S!O&'&!$-&+($?$&-!)+-!()DD&-!$+!(;&!?$'%(!
O&&V!*?!T9+&!JKLL!9%$+D!B>)%($=!%>$B[*+!()D%!B'$+(&-!O$(;!9+$C9&!$-&+($?$=)($*+!
+9:.&'%!Z5&]B)V!3+=RS!G')+V>$+!0C9)'&S!2I\R!5;&!(O*!(/B&%!*?!()DD&-!($BO*':[
$+W9'&-!9B'$D;(%!O&'&!)BB'*]$:)(&>/!&@&+>/!%B)=&-!)>*+D!(;&!&+($'&!>&+D(;!*?!&)=;!
(')+%&=(R!"!(*()>!*?!eK!?>*O&'$+D!)+-!eK!@&D&()($@&!9B'$D;(%!O&'&!()DD&-!?*'!&)=;!
=9>($@)'!ZLd!B&'!(')+%&=(!r!g!(')+%&=(%\R!#)()!*+!(;&!B'&%&+=&!*'!).%&+=&!*?!%$-&[
%;**(%!O&'&!=*>>&=(&-!?'*:!)>>!(;&!()DD&-!9B'$D;(%!$+!0&B(&:.&'!JKLLR!!
5IAIBSG@G@H*GSQNFA*CK*KLCMIB*OED*BISCUNL*C@*ARI*QBCDEFAGC@*CK*TGDI>TRCCAT*G@*
AGQMCBS*G@VEBID*KLCMIBG@H*EQBGHRAT*41GILD*)AEDP*W8*
!E+&!B>*(!Z%&B)')(&!?'*:!?$&>-!%(9-/!L\!$+!&)=;!*?!(;&!=9>($@)'!.&-%!O)%!9%&-R!
G*'!&)=;!(')+%&=(!$+!.*(;!(;&!=9>($@)'!B>*(%S!gd!?>*O&'$+D!9B'$D;(%!O$(;!@$%$.>&!
($BO*':!?&&-$+D!$+W9'/!%$D+%!Z=9BB$+D!*?!(&':$+)>!>&)@&%!O$(;!-&)-!)B$=)>!:&'$%(&:!
)+-!=;)')=(&'$%($=!%=)'!($%%9&\!O&'&!$-&+($?$&-!)+-!()DD&-!$+!(;&!?$'%(!O&&V!*?!T9+&!
JKLL!9%$+D!B>)%($=!%>$B[*+!()D%!B'$+(&-!O$(;!9+$C9&!$-&+($?$=)($*+!+9:.&'%!Z5&]B)V!
3+=RS!G')+V>$+!0C9)'&S!2I\R!5;&!()DD&-!9B'$D;(%!O&'&!)BB'*]$:)(&>/!&@&+>/!%B)=&-!
)>*+D!(;&!&+($'&!>&+D(;!*?!&)=;!(')+%&=(R!"!(*()>!*?!L^K!$+-$@$-9)>!?>*O&'$+D!9B'$D;(%!
O)%!()DD&-!?*'!&)=;!=9>($@)'!Zgd!B&'!(')+%&=(!r!g!(')+%&=(%\R!2*+&!*?!(;&!?>*O&'!.9-%!
!
^d!
O&'&!*B&+!)(!(;&!($:&!*?!()DD$+D!$+!)+/!*?!(;&!9B'$D;(%R!G*'!&)=;!(')+%&=(!$+!.*(;!*?!
(;&!=9>($@)'!B>*(%S!(;&!()DD&-!9B'$D;(%!O&'&!)%%$D+&-!')+-*:>/!(*!*+&!*?!(;&!
?*>>*O$+D!(;'&&!('&)(:&+(%!ZLd!B&'!('&)(:&+(\P!$\!)>>!(;&!?>*O&'!.9-%!'&:*@&-!?'*:!
(;&!9B'$D;(S!$$\!)>>!(;&!?>*O&'!.9-%!'&:*@&-!?'*:!(;&!9B'$D;(!&]=&B(!*+&!Z)(!(;&!.)%&!
*?!JKLL!%&)%*+!D'*O(;!`!(;&!?$'%(!?*':&-!?>*O&'!.9-!*?!)+!9B'$D;(\S!)+-!$$$\!)>>!(;&!
?>*O&'!.9-%!>&?(!$+()=(R!"!')o*'!.>)-&!O)%!9%&-!(*!&]=$%&!(;&!?>*O&'!.9-%R!U*(;!?>*O&'!
'&:*@)>!('&)(:&+(%!O&'&!)BB>$&-!$+!(;&!?$'%(!O&&V!*?!T9+&!ZJKLL\R!#)()!*+!B'&%&+=&!
*'!).%&+=&!*?!%$-&[%;**(%!O&'&!=*>>&=(&-!?'*:!)>>!(;&!()DD&-!9B'$D;(%!$+!0&B(&:.&'!
JKLLR!
5IAIBSG@G@H*GSQNFA*CK*KLCMIB*OED*BISCUNL*C@*ARI*QBCDEFAGC@*CK*TGDI>TRCCAT*G@*
AGQMCBS*G@VEBID*KLCMIBG@H*EQBGHRAT*4-BII@RCETI*)AEDP8*
!5;&!D'&&+;*9%&!%(9-$&%!O&'&!=)''$&-!*9(!O$(;!U&+!4&)'S!H*O&%S!)+-!0(&@&+%R!
1B'$D;(%!*?!U&+!4&)'!O$(;!@$%$.>&!%$D+%!*?!($BO*':!?&&-$+D!$+W9'/!O&'&!=*>>&=(&-!
?'*:!)+!&%().>$%;&-!@)'$&(/!B>*(!)(!(;&!1+$@&'%$(/!*?!A)%%)=;9%&((%!F')+.&''/!
0()($*+!Z6)%(!7)'&;):S!A"h!gLsgetKKRd^u!2!fKsgKtKfR_Ku!7\!$+!(;&!?$'%(!O&&V!*?!
T9+&!JKLLR!0$:$>)'>/S!($BO*':[$+W9'&-!9B'$D;(%!*?!H*O&%!)+-!0(&@&+%!O&'&!=*>>&=(&-!
?'*:!(;&!=*::&'=$)>!?)':!9%&-!$+!G$&>-!0(9-/!LR!3+!)--$($*+!(*!=9''&+(!%&)%*+!
D'*O(;S!&)=;!;)'@&%(&-!9B'$D;(!;)-!qM!=:!*?!B'&@$*9%!%&)%*+!D'*O(;R!G*'!&)=;!
=9>($@)'S!gK!9B'$D;(%!O&'&!B>)+(&-!$+-$@$-9)>>/!$+!LKRJ!=:!%C9)'&!D'*O$+D!B*(%!
Z<'$??$+!<'&&+;*9%&!)+-!29'%&'/!09BB>$&%S!5&OV%.9'/S!A"\!?$>>&-!O$(;!)!MPL!Z%)+-P!
B&)(\!:$](9'&R!E+!(;&!?*>>*O$+D!-)/S!JK!9B'$D;(%!O&'&!%&>&=(&-!')+-*:>/!)+-!)>>!(;&!
?>*O&'!.9-%!O&'&!&]=$%&-!9%$+D!)!')o*'!.>)-&R!5;&!'&:)$+$+D!JK!9B'$D;(%!O$(;!$+()=(!
!
^e!
?>*O&'!.9-%!%&'@&-!)%!=*+('*>%R!F')+.&''/!@$+&%!)'&!'*9($+&>/!B'*B)D)(&-!?'*:!?$&>-[
;)'@&%(&-!9B'$D;(%!)+-!?*':!'**(%!'&)-$>/!9B*+!B>)+($+DR!5;&!B>)+(%!O&'&!
:)$+()$+&-!9+-&'!D'&&+;*9%&!=*+-$($*+%!)+-!*+=&!%($D:)%!.&=):&!&]B*%&-S!)>>!
?>*O&'%!O&'&!;)+-[B*>>$+)(&-!Z0)'')=$+*!)+-!Y*'%)!L__L\!9%$+D!B*>>&+!=*>>&=(&-!
?'*:!(;&!%):&!B>*(%!)%!(;&!9B'$D;(%R!#)()!*+!(;&!B'&%&+=&!*'!).%&+=&!*?!%$-&[%;**(%!
O&'&!=*>>&=(&-!?'*:!)>>!(;&!B>)+(%!$+!0&B(&:.&'!JKLLR!!
5NAN*N@NLPTGT*
7&!9%&-!0"0!@R!_RM!?*'!)>>!(;&!)+)>/%&%!Z0"0!3+%($(9(&!JKLL\!)+-!%&B)')(&!
)+)>/%&%!O&'&!=)''$&-!*9(!?*'!(;&!-$??&'&+(!=9>($@)'%R!
T*/0"($%9)<2*N&($%9)"%6);&9&3"3$;&)'0($953#)<*()35&)0(*6'-3$*%)*<)#$6&:#5**3#)
"<3&()3$0N*(/)$%U'(,)DO$&26)?3'6,)FIR!G*'!&)=;!*?!(;&!?*9'!(')+%&=(%S!(;&!B'*B*'($*+%!
Z)+-!%()+-)'-!&''*'%\!*?!?>*O&'$+D!)+-!@&D&()($@&!9B'$D;(%!O$(;!%$-&[%;**(%!O&'&!
&%($:)(&-R!5;&%&!&%($:)(&%!O&'&!)@&')D&-!*@&'!(;&!?*9'!(')+%&=(%!(*!-&(&':$+&!(;&!
B'*B*'($*+!)+-!@)'$)+=&!*?!($BO*':[$+W9'&-!9B'$D;(%!O$(;!%$-&[%;**(%!Z?>*O&'$+D!
)+-!@&D&()($@&\R!5;&!)@&')D&-!&%($:)(&%!O&'&!9%&-!(*!=*:B)'&!B'*-9=($*+!*?!%$-&[
%;**(%!.&(O&&+!($BO*':[$+W9'&-!?>*O&'$+D!)+-!@&D&()($@&!9B'$D;(%!./!=*+%('9=($+D!
)+!*@&')>>!7)>-*vw!(&%(!*?!+*!-$??&'&+=&!Z,8EF!3A4\!(;)(!)==*9+(%!?*'!(;&!@)'$)+=&!
%('9=(9'&R!5;$%!(&%(!)>>*O&-!(')+%&=(%!(*!.&!)!')+-*:!?)=(*'!)+-!)>%*!?*'!(;&!
('&)(:&+(%!.&$+D!.>*=V&-!*+!(')+%&=(%R!
!&3&(/$%$%9)$/0"-3)*<)<2*N&()K'6)(&/*;"2)*%)35&)0(*6'-3$*%)*<)#$6&:#5**3#)$%)
3$0N*(/)$%U'(&6)<2*N&($%9)'0($953#)ZO$&26)?3'6,)G\R!G*'!&)=;!*?!(;&!?*9'!(')+%&=(%S!(;&!
B'*B*'($*+%!Z)+-!%()+-)'-!&''*'%\!*?!(;&!9B'$D;(%!O$(;!%$-&[%;**(%!$+!(;&!(;'&&!
!
^f!
('&)(:&+(%!O&'&!&%($:)(&-R!5;&%&!&%($:)(&%!O&'&!)@&')D&-!*@&'!(;&!?*9'!(')+%&=(%!
(*!-&(&':$+&!(;&!B'*B*'($*+!)+-!@)'$)+=&!?*'!&)=;!*?!(;&!(;'&&!('&)(:&+(%!)+-!
9($>$o&-!(*!=*+%('9=(!)+!*@&')>>!7)>-*vw!(&%(!*?!+*!-$??&'&+=&!):*+D!(;&!(;'&&!
('&)(:&+(%!Z,8EF!3A4\!(;)(!)==*9+(%!?*'!(;&!@)'$)+=&!%('9=(9'&R!5;$%!(&%(!)>>*O&-!
(')+%&=(%!(*!.&!)!')+-*:!?)=(*'!)+-!)>%*!?*'!(;&!('&)(:&+(%!.&$+D!.>*=V&-!*+!
(')+%&=(%R!,)$'O$%&!=*:B)'$%*+%!):*+D!(;&!(;'&&!('&)(:&+(%!O&'&!=)''$&-!*9(!9%$+D!
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!Bast, M., and R. J Reader. 2003. Regrowth response of young black spruce (Picea
mariana) trees to meristem removal and resource addition. Can. J. Botany 81:
956-963.
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1+$@&'%$(/!*?!F)>$?*'+$)!,'&%%S!U&'V&>&/!)+-!4*%!"+D&>&%S!F"R!!!Crawley, M. J. 1997. Plant – herbivore dynamics, pp. 401-474. In M. J. Crawley (ed.),
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Scammell, H. B. 1917. Cranberry insect problems and suggestions for solving them.
United States Department of Agriculture Farmers’ Bulletin 860: 14-16.
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!Tiffin, P. 2000. Mechanisms of tolerance to herbivore damage: what do we know? Evol.
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!Tolvanen, A., and K. Laine. 1997. Effects of reproduction and artificial herbivory on
vegetative growth and resource levels in deciduous and evergreen dwarf shrubs.
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