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TropicalNativeTreePlantations:
AnInvestigationofHerbivoreDamageandSpecies
SelectioninCentralPanama
Gillian Paul, MFS 2010
Introduction
0U [OL 9LW\ISPJ VM 7HUHTH HYV\UK VM
MVYLZ[Z ^LYL SVZ[ IL[^LLU HUK SHYNLS`
K\L[VJVU]LYZPVUPU[VHNYPJ\S[\YLHUKWHZ[\YLSHUKZ
9LJLU[S`THU`VM[OVZLHNYPJ\S[\YHSHUKNYHaPUNHYLHZ
OH]L ILLU HIHUKVULK HSSV^PUN [OL VWWVY[\UP[` MVY
UH[\YHSYLNLULYH[PVUHUKYLMVYLZ[H[PVU6M7HUHTHUPHU
SHUKZ JVU]LY[LK [V [PTILY WSHU[H[PVUZ HU HSHYTPUN
VMZ[HUKZHYLWSHU[LK^P[OL_V[PJ[YLLZ*YH]LUL[
al. 2007). In Latin America as a whole, reforestation
LMMVY[Z MVJ\Z VU [OL WSHU[PUN VM ^LSSRUV^U L_V[PJ
trees of Pinus, Eucalyptus, and Tectona species
,]HUZ HUK ;\YUI\SS ;OLZL L_V[PJ [YLLZ HYL
favorable to plantation foresters because of the ease
VMTHUHNPUNHZ[HUK^P[OZPTWSPÄLKU\YZLY`WYHJ[PJLZ
THUHNLTLU[[LJOUPX\LZHUKOHY]LZ[PUN[PTLZ7PV[[V
(S[OV\NOVM[LULJVUVTPJHSS`WYVÄ[HISL[OLZL
Z`Z[LTZVMMLYML^LYLJVSVNPJHSILULÄ[ZVMYLZ[VYH[PVU
MVYVYNHUPZTZ[OH[KLWLUKVUYLZV\YJLZZ\JOHZMY\P[
nectar, and habitat structure provided by native
species (Hartley 2002).
early in plantation establishment.
6UL IV[[SLULJR [V WSHU[H[PVU MVYLZ[Y` HZ H
means of reforestation can be reduced survivability
HUKWYVK\J[P]P[`VM[OL[YLLZK\L[VOLYIP]VY`5H[P]L
timber trees may be more susceptible to the native
PUZLJ[ OLYIP]VYLZ PU H YLNPVU (KKP[PVUHSS` \USPRL
the detailed recorded literature about pest control
practices of pine, eucalyptus, and teak monocultures,
[OLYLOH]LILLUML^LYYLZLHYJOZ[\KPLZPU]LZ[PNH[PUN
WLZ[JVU[YVSMVYUH[P]L[PTILYWSHU[H[PVUZ)\[[LYÄLSK
(S[OV\NO L_V[PJ ZWLJPLZ JHU OH]L KHTHNPUN
insect outbreaks, native species plantations may be
hard to control for herbivory because there are much
fewer resources available on pest control practices to
N\PKLSHUKV^ULYZHUKMVYLZ[WYHJ[P[PVULYZ
Tree species selection, which can be different
KLWLUKPUN VU [OL WHY[PJ\SHY YLNPVU VM [OL ^VYSK PZ
SPRLS`[VOH]LHZ[YVUNPTWHJ[VU[OL]\SULYHIPSP[`VM
[OLWSHU[H[PVU[VPUZLJ[KHTHNLHUKHMMLJ[[OLZ\JJLZZ
In comparison, native species plantations have of the overall reforestation effort. Some tree species
been demonstrated to better promote biodiversity and have evolved different mechanical and chemical
PTWYV]L ZP[L X\HSP[` 3\NV 0U 7HUHTH HSVUL JOHYHJ[LYPZ[PJZ [V TPUPTPaL PUZLJ[ JVUZ\TW[PVU
there are more than 100 native species valued for [OLYLI` YLK\JPUN [OL Z[YLZZ VM OLYIP]VY` VU [OL
timber and other uses (Wishnie et al. 2007). Scientists photosynthetic abilities of plants (Coley and Barone
HUK YLZLHYJO PUZ[P[\[PVUZ OH]L ILLU PUJYLHZPUN ">OP[LHUK:JV[[9LZLHYJOVU[OLNYV\UK
their study of mixed native species plantations, but is needed to evaluate how adaptations of different
those plantations are still rarely used as a means of species, in the form of leaf traits, affect the resistance
MVYLZ[ YLZ[VYH[PVU -VY T` YLZLHYJO 0 ^HZ L_JP[LK [V VM[OVZL[YLLZ[VPUZLJ[KHTHNL
PU]LZ[PNH[LUH[P]LZWLJPLZWSHU[H[PVUZPU3H[PU(TLYPJH
and evaluate how the selection by humans of tree -VY T` YLZLHYJO 0 HUHS`aLK [OL SLHM HYLH
ZWLJPLZMVYWSHU[PUNYLSH[LZ[V]\SULYHIPSP[`[VKHTHNL KHTHNL MVY UH[P]L [PTILY]HS\L [YLL ZWLJPLZ PU
7HUHTH! Terminalia amazonia, Dalbergia retusa,
Pachira quinata, Tabebuia rosea, and Anacardium
excelsum. These species are valued for their timber
Gillian Paul grew up in Texas and Virginia. She has
PU7HUHTHHUKHYLYLWYLZLU[H[P]LVMHKP]LYZLYHUNL
been working locally and internationally conducting VM MHTPSPLZ SPNO[ [VSLYHUJL HUK NYV^[O OHIP[Z
research in forest health and participating in envi- The objective of this study is to identify if there are
ronmental outreach initiatives. She graduated with differences in the foliar herbivory between these
a Master of Forest Science degree from Yale F&ES native tree species and evaluate how well measured
and hopes to participate in the implementation of leaf traits relate to the susceptibility to insect
conservation projects that advance both scientific KHTHNL )` L]HS\H[PUN [OL SLHM HKHW[H[PVUZ VM [OL
understanding and stewardship of forest resources. native tree species for resistance to herbivory, I seek
22 Volume 29, 2010
Tropical Native Tree Plantations in Central Panama
Photo 1. The rolling hills of the Agua
Salud Plantations of
native tree species in
central Panama
to draw comparisons on the relationship between
the protective mechanisms and the extent of insect
defoliation.
[OLWSHU[;OPZ^HZULJLZZHY`ILJH\ZL[OLZHTWSLZPaL
ULLKLK[VNL[HUHJJ\YH[LL]HS\H[PVUVMSLHMKHTHNL
on the plant would have necessitated the collection
VM H SHYNL WVY[PVU VM [OL WOV[VZ`U[OL[PJ HYLH VM
[OLZLZ`V\UN[YLLZHWSPUNZ(SZVMVYTLHZ\YLTLU[VM
Methods
OLYIP]VY`0SVVRLKH[KPMMLYLU[JH[LNVYPLZVMOLYIP]VY`
My research was conducted in central JOL^PUN ZRLSL[VUPaPUN TPUPUN ZOLS[LY YVSSPUN
7HUHTHH[[OLZP[LVM[OL(N\H:HS\K7YVQLJ[HWYVQLJ[ HUK NHSS YVSSPUN [OH[ NP]L PUZPNO[ PU[V [OL M\UJ[PVUHS
formed as a partnership by the Smithsonian Institution MLLKPUNNYV\WZVMPUZLJ[ZJH\ZPUN[OLSLHMKHTHNL
and Panama Canal Watershed Authority to develop
]PHISLZ[YH[LNPLZMVY[OLYLZ[VYH[PVUVMMVYLZ[SHUKHUK
The second half of my study consisted of
LJVZ`Z[LT ZLY]PJLZ VU KLNYHKLK SHUKZ ;OL [OL JVSSLJ[PVU VM SLH]LZ MVY SLHM [YHP[ HUHS`ZPZ 6U
LZ[HISPZOTLU[VM[OL(N\H:HS\KWSHU[H[PVUWYV]PKLK separate trees in each plot, four leaves from each
an excellent opportunity to compare insect defoliation tree were collected and stored in a cooler to retain
VM SLH]LZ HTVUN UH[P]L [YLL ZWLJPLZ WSHU[LK PU SLHM TVPZ[\YL >P[OPU OV\YZ MYVT JVSSLJ[PVU 0
hectares of previously abandoned pasture. The land [LZ[LK [OL SLH]LZ MVY SLHM [V\NOULZZ H TLHZ\YLTLU[
use history of the land consists of corn, rice, and other VM W\UJ[\YL Z[YLUN[O [OH[ NP]LZ HU PUKPJH[PVU VM [OL
JYVWJ\S[P]H[PVUHZ^LSSHZWHZ[\YLNYHaPUN\U[PSYLJLU[ relative ease at which an insect can penetrate the
SHUKHIHUKVUTLU[#`LHYZHNV
surface of a leaf for consumption.
-YVT4H`\U[PS(\N\Z[ 0JVSSLJ[LKKH[H
VU[OLOLYIP]VY`HUKSLHM[YHP[ZVM`V\UN[YLLZHWSPUNZ
of the timber trees T. amazonia, D. retusa, P. quinata,
T. rosea, and A. excelsum(TVUN[OLOLJ[HYLZVM
TP_LKUH[P]LZWLJPLZVU[OLSHUKZJHWL0^VYRLKPU
I`[YLLZPUNSLZWLJPLZWSV[Z-VY[OLÄYZ[OHSMVMT`
Z[\K`OLYIP]VY`VUH[YLLSL]LS^HZLZ[PTH[LK[OYV\NO
HJVTIPUH[PVUVM[^VMHJ[VYZKPNP[HSHUHS`ZPZVMHYLH
KHTHNLKWLYSLHMHUKHULZ[PTH[LVMSLH]LZKHTHNLK
WLY WSHU[ 0UZ[LHK VM JVSSLJ[PUN SLH]LZ MVY OLYIP]VY`
HUHS`ZPZT`TL[OVKVMKPNP[HSS`TLHZ\YPUNOLYIP]VYL
KHTHNL^HZ\UPX\LS`KLZPNULK[VSL[HSLHMYLTHPUVU
5L_[0TLHZ\YLK[OLMYLZOSLHMHYLH\ZPUNH
KPNP[HSZJHUULY3LHMZHTWSLZ^LYLKYPLKHUK^LPNOLK
[V JHSJ\SH[L ZWLJPÄJ SLHM HYLH :3( :3( MYLZO SLHM
area divided by dry leaf mass (cm2N-1), is known to
KLJYLHZL ^P[O SLHM HNL HZ [OL SLH]LZ OHYKLU K\L [V
[OLOPNOLYKLUZP[`[PZZ\L[OPJRLYSHTPUHHUKZ[YVUNLY
veins associated with low-SLA leaves (Moles and
>LZ[VI` -PUHSS` SLH]LZ ^LYL NYV\UK HUK
TLHZ\YLK MVY JHYIVU HUK UP[YVNLU ]HS\LZ I` TLHUZ
VM [OLYTHS JVTI\Z[PVU 3HYNLY UP[YVNLU HUK ZTHSSLY
*HYIVU!5P[YVNLUYH[PVZZ\NNLZ[TVYLWHSH[HIPSP[`HUK
U\[YP[PVUHS]HS\LVM[OLSLHM[VMLLKPUNPUZLJ[Z
Tropical Resources Bulletin 23
Gillian Paul
Results and Discussion
TH`ILJVUMV\UKPUN[OL[YLUKZPU[OLSLHM[YHP[Z(SZV
MVYM\[\YLZ[\K`L]HS\H[PUN[OLWYLZLUJLVMZLJVUKHY`
6]LYHSS[OLOLYIP]VYLKHTHNL^HZSLZZ[OHU defense mechanisms, or toxic compounds in the
VM[OLSLHMHYLHWLYWSV[MVYLHJOZWLJPLZ;OLZL SLH]LZ ^PSS IL PTWVY[HU[ PU \UKLYZ[HUKPUN [OL SV^
Ä]L ZWLJPLZ ^LYL VYPNPUHSS` JOVZLU ILJH\ZL VM [OLPY levels of herbivory.
]PNVY PU UH[P]L ZWLJPLZ WSHU[H[PVUZ PU V[OLY HYLHZ VM
7HUHTH ]HU )Y\LNLS PU WYLZZ ,]LU [OV\NO SLHM 6UL PU[LYLZ[PUN ÄUKPUN WLYOHWZ TVYL
KHTHNL ^HZ SV^ [OLYL ^LYL ZPNUPÄJHU[ KPMMLYLUJLZ important than leaf traits I measured, was that insect
HTVUN[OLZWLJPLZ-PN;OLKHTHNL[VA. excelsum KHTHNL YLSH[LK [V [OL WOLUVSVN` HUK Z\JJLZZPVUHS
and T. rosea ^HZ ZPNUPÄJHU[S` OPNOLY [OHU [OL V[OLY state in which the species is found in the forest.
ZWLJPLZ^OPSLKHTHNL[VD. retusa and P. quinata was Anacardium excelsum was the only plant with
ZPNUPÄJHU[S`SV^LY[OHU[OLV[OLYZWLJPLZAnacardium L]LYNYLLU SLH]LZ @V\UN SLH]LZ OHK SV^ SL]LSZ VM
excelsum and T. rosea also had more diversity in the OLYIP]VY` HUK OPNOLY [V\NOULZZ ]HS\LZ [OH[ V[OLY
[`WLZVMPUZLJ[KHTHNL[OHUV[OLYZWLJPLZ
species. Mature leaves of this species demonstrated
[OL OPNOLZ[ OLYIP]VY` HUK ]HYPL[` VM KHTHNL [`WLZ
The correlations between leaf traits and ^OPSL HSZV L_OPIP[PUN [OL OPNOLZ[ [V\NOULZZ HUK
OLYIP]VY` PZ KLWPJ[LK PU ÄN\YL *VU[YHY` [V T` [OL SV^LZ[ UP[YVNLU JVUJLU[YH[PVU 3P[LYH[\YL VU A.
O`WV[OLZPZ [OL ZWLJPLZ ^P[O OPNOLY OLYIP]VY` excelsum Z\NNLZ[Z [OH[ SLH]LZ VM [OPZ ZWLJPLZ OH]L
KLTVUZ[YH[LK OPNOLY SLHM [V\NOULZZ HUK SV^LY :3( OPNOLY WOLUVSZ H [`WL VM JOLTPJHS KLMLUZL [OHU
/V^L]LY SLHM [V\NOULZZ ^HZ UV[ MV\UK [V IL H many other species in Panama (Coley et al. 2002).
ZPNUPÄJHU[WYLKPJ[VYVMPUZLJ[KHTHNLPU[OPZZ[\K`0[ 0[ PZ WVZZPISL [OH[ [OL OPNOLY [V\NOULZZ HUK SV^LY
PZWVZZPISL[OH[[OL[V\NOULZZVM[OLZLZWLJPLZH[[OLPY UP[YVNLUJVU[LU[WYV[LJ[SLH]LZ^OPSL[OL`HYL`V\UN
J\YYLU[ HNL PZ UV[ OPNO LUV\NO [V KL[LY OLYIP]VYLZ HUK\UOHYKLULK;OLOPNOLYYH[LZVMOLYIP]VY`JV\SK
HUK[OLYLI`PZ\UYLSH[LK[V[OLPYWHSH[HIPSP[`;OL*!5 ILILJH\ZL[OPZL]LYNYLLUZWLJPLZOHZSLH]LZL_WVZLK
YH[PVZKLTVUZ[YH[LKHZPNUPÄJHU[WVZP[P]LJVYYLSH[PVU HUKH]HPSHISLMVYLH[PUNMVYSVUNLY[PTL[OHUSLH]LZVM
HUK UP[YVNLU ]HS\LZ L_OPIP[LK H ZPNUPÄJHU[ ULNH[P]L other species.
JVYYLSH[PVU^P[OOLYIP]VY`:PTPSHY[VSLHM[V\NOULZZ
[OPZ ÄUKPUN ¶ [OH[ SLH]LZ ^P[O OPNOLY OLYIP]VY` OHK
Herbivory in the other species relate to the
SV^LYSLHMUP[YVNLU¶^HZKPMMLYLU[MYVTT`WYLKPJ[PVUZ successional state in which those trees are found
The small differences in herbivory between species in the wild. Pachira quinata is a mid-successional
Figure 1. Total Herbivory (standard error bars) for mature and expanding leaves
24 Volume 29, 2010
Tropical Native Tree Plantations in Central Panama
ZWLJPLZNYV^PUN[VJHUVW`KVTPUHUJLPU[OLWLYPVK
between the initial forest and the later mature forest.
The herbivory of P. quinata was relatively low in
JVTWHYPZVU ^P[O [OL V[OLY ZWLJPLZ L]LU [OV\NO P[
L_OPIP[LK[OLSV^LZ[SLHM[V\NOULZZPUIV[OL_WHUKPUN
and mature leaves. More research is needed to
determine the chemical defenses of P. quinata beyond
SLHMUP[YVNLU
;OLOPNOOLYIP]VY`]HS\LZMVYT. rosea and T.
amazonia may be related to the early-successional
OHIP[H[ VM [OVZL ZWLJPLZ )LPUN LHYS` Z\JJLZZPVUHS
[OL` SPRLS` PU]LZ[ [OLPY YLZV\YJLZ PU X\PJR NYV^[O H[
[OLL_WLUZLVMWYV[LJ[PVUHNHPUZ[OLYIP]VYLZ)LJH\ZL
VM [OL OPNO [V\NOULZZ HUK SV^ :3( L_OPIP[LK PU
these species, they may depend primarily on these
TLJOHUPJHS KLMLUZLZ TVYL [OHU LULYN` PU[LUZP]L
secondary chemicals. There were two plots of T. rosea
PU ^OPJO OPNO HTV\U[Z VM YVSSPUN OLYIP]VY` ^LYL
MV\UKVU`V\UNL_WHUKPUNSLH]LZ;OLZLWSV[Z^LYL
not in spatial proximity on the landscape. Therefore,
[OLWYLZLUJLVMYVSSPUNOLYIP]VY`PZSPRLS`K\L[V[^V
ZLWHYH[L PUMLZ[H[PVUZ VM YVSSPUN PUZLJ[Z ZWLJPÄJHSS`
H[[YHJ[LK[V`V\UNT. rosea leaves.
Dalbergia retusa is a late successional,
SLN\TPUV\Z ZWLJPLZ 0U [OL SLHM [YHP[ HUHS`ZPZ D.
retusaL_OPIP[LK[OLOPNOLZ[UP[YVNLUJVU[LU[I\[HSZV
maintained the lowest herbivory in mature leaves.
(S[OV\NOOPNOLYUP[YVNLUJVUJLU[YH[PVUZJHUILTVYL
nutritious to animals, this species clearly demonstrated
H OPNOLY YLZPZ[HUJL [V PUZLJ[ KHTHNL HM[LY SLHM
TH[\YH[PVU3H[LZ\JJLZZPVUHS[YLLZVM[LUNYV^PU[OL
ZOHK`\UKLYZ[VY`^OPSLMHZ[LYNYV^PUNZWLJPLZL_OPIP[
dominance early in forest succession. Their position
in the shady understory can be a favorable habitat
[V OLYIP]VYLZ" [OLYLMVYL P[ PZ \UKLYZ[HUKHISL [OH[D.
retusa would be more adapted to resist herbivory. As
SVUN HZ [OPZ ZOHKL [VSLYHU[ ZWLJPLZ HSZV NYV^Z ^LSS
PU[OLOPNOSPNO[LU]PYVUTLU[VMHIHUKVULKWHZ[\YL
then it is a wise choice for use in a native species
plantation.
Tropical Resources Bulletin 25
Gillian Paul
Conclusions
This research is meant to serve as a resource
MVYMVYLZ[THUHNLYZHUKWYHJ[P[PVULYZJOVVZPUNZWLJPLZ
for tropical forest restoration and plantations. Tropical
forest plantations are important for timber production,
JSLHU^H[LYZ\WWS`JHYIVUZLX\LZ[YH[PVUHUKOHIP[H[
MVYIPVKP]LYZP[`5H[P]LZWLJPLZWSHU[H[PVUZJHUZLY]L
HZ JH[HS`Z[Z MVY MVYLZ[ YLOHIPSP[H[PVU PU KLNYHKLK
[YVWPJHS HYLHZ ^OPSL WYV]PKPUN LU]PYVUTLU[HS HUK
LJVUVTPJILULÄ[Z[V[OLYLNPVU7HYYV[[HL[HS 0UZ[LHK VM YLS`PUN VU H ML^ ^LSS Z[\KPLK L_V[PJ [YLL
ZWLJPLZ MVYLZ[ THUHNLYZ ^VYSK^PKL JHU \ZL [YLL
JOHYHJ[LYPZ[PJZ PU [OLPY JVUZPKLYH[PVUZ MVY ZLSLJ[PUN
[YLLZUH[P]L[V[OLPYV^UYLNPVU
JOHYHJ[LYPZ[PJZHUKMVSPHYU\[YPLU[JVU[LU[VM
tree species at a wet, fertile site and a dry, infertile
ZP[LPU7HUHTH-VYLZ[,JVSVN`HUK4HUHNLTLU[
!
,]HUZ1HUK1>;\YUI\SS7SHU[H[PVU-VYLZ[Y`PU
[OL;YVWPJZYKLK*SHYLUKVU7YLZZ6_MVYK<2
Hartley, M. J. 2002. Rationale and methods for conservPUNIPVKP]LYZP[`PUWSHU[H[PVUMVYLZ[Z-VYLZ[,JVSVN`
HUK4HUHNLTLU[! 3HTI+ 3HYNLZJHSLLJVSVNPJHSYLZ[VYH[PVUVM
KLNYHKLK[YVWPJHSMVYLZ[SHUKZ![OLWV[LU[PHSYVSLVM
[PTILYWSHU[H[PVUZ9LZ[VYH[PVU,JVSVN`! 3\NV(, ;OLHWWHYLU[WHYHKV_VMYLLZ[HISPZOPUN
ZWLJPLZYPJOULZZVUKLNYHKLKSHUKZ^P[O[YLLTVUV
0U [OLPY ÄYZ[ ^L[ ZLHZVU PU [OL NYV\UK [OL J\S[\YLZ-VYLZ[,JVSVN`HUK4HUHNLTLU[ ! native species in this study demonstrated low levels
VMOLYIP]VYLKHTHNL6MJV\YZLM\Y[OLYYLZLHYJO^PSS
ULLK [V KL[LYTPUL PM [OPZ [YLUK JVU[PU\LZ" OV^L]LY
[OPZ ÄUKPUN PZ H NVVK ZPNU MVY [OL ]PHIPSP[` VM UH[P]L
ZWLJPLZWSHU[H[PVUZ-VYZLSLJ[PVUVMZWLJPLZJOVVZPUN
SH[L Z\JJLZZPVUHS ZWLJPLZ [OH[ HYL HSZV SPNO[[VSLYHU[
such as D. retusa, may help reduce the palatability
VM[OLPY[YLLZ[VMLLKPUNPUZLJ[Z-\Y[OLYPU]LZ[PNH[PVU
is needed to evaluate chemical defenses in these
ZWLJPLZIL`VUKSLHMUP[YVNLU
It is my hope that the results of my study can
inspire the use of native species for other plantation
WYVQLJ[Z HUK LUJV\YHNL M\Y[OLY YLZLHYJO HPTLK
H[ WYV]PKPUN MVYLZ[ THUHNLYZ ^P[O PUZPNO[ PU[V [OL
LJVSVNPJHS JVUZPKLYH[PVUZ MVY ZLSLJ[PUN WSHU[H[PVU
species most likely to survive early in plantation
establishment.
9LMLYLUJLZ
)\[[LYÄLSK97 7YVTV[PUNIPVKP]LYZP[`!HK]HUJLZPU
L]HS\H[PUNUH[P]LZWLJPLZMVYYLMVYLZ[H[PVU-VYLZ[
,JVSVN`HUK4HUHNLTLU[!
*VSL`7+HUK1()HYVUL /LYIP]VY`HUKWSHU[
defenses in tropical forests. Annual Review of
,JVSVN`HUK:`Z[LTH[PJZ!
*VSL`7+44HZZH*,3V]LSVJRHUK2>PU[LY
,MMLJ[ZVMLSL]H[LK*6VUMVSPHYJOLTPZ[Y`VM
ZHWSPUNZVMUPULZWLJPLZVM[YVWPJHS[YLL6LJVSVNPH
! *YH]LU++)YHKLU4:(ZO[VU.7)LYS`U4
Wishnie, and D. Dent. 2007. Between and withinZP[LJVTWHYPZVUZVMZ[Y\J[\YHSHUKWO`ZPVSVNPJHS
26 Volume 29, 2010
Moles A., and M. Westoby 2000. Do small leaves expand
MHZ[LY[OHUSHYNLSLH]LZHUKKVZOVY[LYL_WHUZPVU
[PTLZYLK\JLOLYIP]VYLKHTHNL&6PRVZ !
¶
7HYYV[[H1(1>;\YUI\SSHUK51VULZ 0U[YV
K\J[PVUJH[HS`aPUNUH[P]LMVYLZ[YLNLULYH[PVUVU
KLNYHKLK[YVWPJHSSHUKZ-VYLZ[,JVSVN`HUK
4HUHNLTLU[ !
7PV[[V+(TL[HHUHS`ZPZJVTWHYPUN[YLLNYV^[OPU
TVUVJ\S[\YLZHUKTP_LKWSHU[H[PVUZ-VYLZ[,JVSVN`
HUK4HUHNLTLU[!
>OP[L1+HUK5(:JV[[:WLJPÄJSLHMHYLHHUK
UP[YVNLUKPZ[YPI\[PVUPU5L^ALHSHUKMVYLZ[Z!ZWL
JPLZPUKLWLUKLU[S`YLZWVUK[VPU[LYJLW[LKSPNO[ -VYLZ[,JVSVN`HUK4HUHNLTLU[! >PZOUPL4/+/+LU[,4HYPZJHS1+LHNV5
Cedeno, D. Ibarra, R. Condit, and P. M. S. Ashton.
2007. Initial performance and reforestation potential
VM[YVWPJHS[YLLZWLJPLZWSHU[LKHJYVZZHWYLJPW
P[H[PVUNYHKPLU[PU[OL9LW\ISPJVM7HUHTH-VYLZ[
,JVSVN`HUK4HUHNLTLU[!