Integrating genetic and silvicultural strategies to minimize abiotic and biotic constraints in Brazilian

22 Pages·2013·1 MB·English

ForestEcologyandManagementxxx(2013)xxx–xxx
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Forest Ecology and Management
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Integrating genetic and silvicultural strategies to minimize abiotic and biotic
constraints in Brazilian eucalypt plantations
José Leonardo de Moraes Gonçalvesa,⇑, Clayton Alcarde Alvaresb, Antonio Rioyei Higac,
Luciana Duque Silvaa, Acelino Couto Alfenasd, James Stahle, Silvio Frosini de Barros Ferraza,
Walter de Paula Limaa, Pedro Henrique Santin Brancaliona, Ayeska Hubnera, Jean-Pierre Daniel Bouilletf,a,
Jean-Paul Laclauf,a, Yann Nouvellonf,g, Daniel Epronf,h
aUSP,UniversidadedeSãoPaulo,ESALQ,DepartamentodeCiênciasFlorestais,CEP13418-900Piracicaba,SP,Brazil
bInstitutodePesquisaseEstudosFlorestais(IPEF),Piracicaba,SP,Brazil
cUniversidadeFederaldoParaná,Curitiba,PR,Brazil
dUniversidadeFederaldeViçosa,MG,Brazil
eKlabin,TelêmacoBorba,PR,Brazil
fCIRAD,UMR111,EcologieFonctionnelle&BiogéochimiedesSols&Agro-écosystèmes,F-34060Montpellier,France
gUSP,UniversidadedeSãoPaulo,DepartamentodeCiênciasAtmosféricas,IAG,CEP05508-900SãoPaulo,Brazil
hUniversitédeLorraine,UMRINRA-UL1137,EcologieetEcophysiologieForestières,FacultédesSciences,Vandoeuvret-les-Nancy,France
a r t i c l e i n f o a b s t r a c t
Articlehistory: OrganizedforestryinBrazilbeganinthelate1960s,stimulatedbyagovernmentpolicywhichsubsidized
Availableonlinexxxx afforestationprogramsfrom1967to1989todevelopaninternationally-competitivewood-basedindus-
try,managedbytheprivatesector.Currently,plantedforestsinBraziltotalabout6.9millionha,from
Keywords: which4.9millionhaisplantedwitheucalypt(around25%ofworldplantation),1.6millionhawithpine,
Geneticadaptation and0.42Mhawithotherspecies.Roundwoodconsumptionofforestplantationstotaled170.1millionm3
Silviculture in2011,eucalyptplantationaccountedfor80.6%ofthistotal.
Forestry
Mosteucalyptplantationsaremanagedinshortrotations(6–8years)andareestablishedinregions
Abiotic
withwater,nutritionalandfroststressesoflowtohighdegrees.Themeanannualincrementis40m3
Biotic
ha(cid:2)1year(cid:2)1roundwood,rangingfrom25to60m3ha(cid:2)1year(cid:2)1dependingonthelevelofenvironmental
Stress
stress.Improvingnaturalresourcesuseefficiencybybreedingandmatchinggenotypestositesandusing
appropriatesitemanagementpracticesisakeychallengetosustainorincreaseproductivity.
Thewiderangeofeucalyptspeciesandhybridswithdifferentclimaticandedaphicsuitabilityassoci-
atedwiththeeasypropagationbyseedsandcloningallowtheadaptationofplantationstovarioustrop-
icalandsubtropicalregionsinBrazil.Thepossibilityofusingeucalyptwoodinarangeofpurposeshasled
largeandsmallenterprisestoestablisheucalyptforestsformultipleuses.Thedesirablecharacteristicsin
associationwiththeaccumulatedknowledgeoneucalyptsilvicultureencouragetheuseofthisgenusin
mostplantations.Themostimportantfactorsintheselectiveprocessforagenotypearewoodcharacter-
istics, productivity level, susceptibility to pests and diseases, drought tolerance, especially in tropical
regions(frostfree),andfrosttoleranceinsubtropicalregions(mostlywithoutwaterdeficit).Inregions
withpronouncedseasonalityandmoderatetolongdroughtperiods,theplantingofhybridgenotypes
predominates,propagatedbycloning.Undersubtropicalconditions,theplantingofsinglespeciespre-
dominates, propagated by seed. Clonal plantations with interspecific hybrids have been fundamental
foreucalyptadaptationinregionsunderwaterandnutritionalstresses.Giventherapidadvancesineuca-
lyptbreeding,regardingadaptationtowaterstressandresistancetodiseasesandpests,andtheadoption
ofclonalpropagationtechniques,genotypesarerapidlybecomingobsoleteandarereplacedbymorepro-
ductiveonesafterharvesting.Thus,thereplantingofcropshasbecomeacommonprocedureafterthe
secondhalfofthe1990sinBrazil.
Thispaperdescribesthebasicrequirementsforintegratinggeneticandsilviculturalstrategiestomin-
imizeabioticandbioticconstraintsineucalyptplantations.
(cid:2)2013ElsevierB.V.Allrightsreserved.
⇑
Correspondingauthor.Tel.:[phone];fax:[phone].
E-mailaddress:[email protected](J.L.M.Gonçalves).
0378-1127/$-seefrontmatter(cid:2)2013ElsevierB.V.Allrightsreserved.
http://dx.doi.org/10.1016/j.foreco.2012.12.030
Pleasecitethisarticleinpressas:Gonçalves,J.L.M.,etal.IntegratinggeneticandsilviculturalstrategiestominimizeabioticandbioticconstraintsinBra-
zilianeucalyptplantations.ForestEcol.Manage.(2013),http://dx.doi.org/10.1016/j.foreco.2012.12.030
2 J.L.M.Gonçalvesetal./ForestEcologyandManagementxxx(2013)xxx–xxx
1.Backgroundofeucalyptplantations studieswerefundedbythePaulistaRailwayCompanyandcoordi-
natedbyEdmundoNavarrodeAndrade.By1919,theseedorchard
Brazilianforestrycanbedividedintotwomainperiods,before ofRioClarohadturnedout123speciesofEucalyptus,whichwere
andafterthe20thcentury.Itwasstartedasnativesilviculturewith studied for seedling production, soil preparation, alignment and
theexperienceofreforestationofTijucaForest(RiodeJaneiro)in spacing of planting, pruning, thinning, harvesting, coppicing, dis-
1862,whichaimedtorestoretheAtlanticForestinsitesformerly eases,pests,productivity,andusesofwood(Andrade,1961).This
used for coffee plantations (Silva, 1870). This was an important phasemarksthebirthofscientificforestryandintensiveforestry
milestonebecauseitwasthefirstsuccessfulreforestationthrough- inBrazil.
outAmericas.Thisperiodwasmarkedbytraditionalforestrythat Untilthe1930s,theplantationshadasmainobjectivestogrow
wasruledbyvaluesofconservationandecology,untilthenprac- upspecieswithpropertimbertosupplythedemandsoffirewood
ticedinthenorthernhemisphere.Onlynativespecieswereused, usedasfuelforlocomotivesandsleepersforrailways,whichwere
regeneration was based on natural methods, where clearcuts in inplainexpansionin Brazil.Theannualplantingofeucalyptwas
largeareas,intensivesitepreparationandapplicationoffertilizer about 1000hayr(cid:2)1 (Andrade, 1923). Some years later, based on
werenot practiced(Ferreira,1989). Inthis period, thefirst batch the results of early experiments comparing native species with
of eucalypt seeds coming from Montevideo, Uruguay arrived in eucalyptscultivatedinJundiaiandRioClaro,itwasconcludedthat
Brazil (Azevedo, 1874). Eucalypt plantations were quickly wide- species of Eucalyptus should be planted on a commercial scale
spread in the states of Rio de Janeiro, Sao Paulo, Parana and Rio (Andrade, 1928, 1936). Shortly afterwards, the planted area in-
Grande do Sul, mainly used as urban forestry and hedgerows for creased to about 25,000hayr(cid:2)1 (Reis, 1948). Since 1950s started
farms. Although the government had encouraged the search for theestablishmentoflargeplantationsfortheproductionofchar-
eucalypt acclimation at that time, there was no success, because coalinSãoPaulo(Alvares,2011).Overthesameperiod,thepulp
theworkswereverywidespreadandnotproperlycoordinated. and paper industry adopted eucalypt as the main source of raw
The first coordinated research studies of eucalypt forestry material(Lemos,2012).
started in early twentieth century, more specifically in 1904, in Itwasfromthe1960sthatBrazilianforestryhadabigboost.Be-
the municipality of Jundiai, and in 1909 in the municipality of tweenthe1960sandthe1980s,theforestrysectorwasencouraged
Rio Claro (Andrade, 1909), both in Sao Paulo state. These early to develop forest plantations by a governmental policy that
granted tax incentives (Law n. 5106, 2 September 1966, giving
tax incentives to afforestation). In the beginning, about
37,000hayr(cid:2)1 were planted with Eucalyptus (Bastos, 1961), and
at the end of period of tax incentives, about 270,000hayr(cid:2)1
(Fig. 1a). Productivity was relatively low ranging from 10 to
30m3ha(cid:2)1year(cid:2)1 (Ferreira, 1992; Campinhos, 1999). High rate
of segregation caused by uncontrolled hybridizations, major
changesin vigor and woodqualityshowed that theseed sources
were not adequate. Consequently, the phenotypic variability of
themost plantedspecies, namelyEucalyptus grandis(over 70% of
theplantedarea),EucalyptusurophyllaandEucalyptussalignawere
verylarge(Ferreira,1992).Tomazello(1976)obtainedremarkable
findingsinidentifyingthehighcankerresistanceatanindividual
level of these species. The author also found that trees with
gummed-bark like (smooth) were the most resistant. These find-
ingsconstitutedoneofthebasicguidelinesfortheclonalmultipli-
cationofsuperiorindividuals(Ferreira,1992).Seekingproductivity
increase and wood quality improvement, new species and new
provenancesofEucalyptus,suchasE.grandis(Coff’sHarbour,Aus-
tralia), and E. urophylla (Timor and Flores, Indonesia) were intro-
duced.TheintroductionofE.grandis(Coff’sHarbour)provenance
occurred after the visit of Professor LD Pryor to Brazil (Pryor,
1971).Atthattime,theecologicalzoningofexoticspecies(Golfari,
1974), the selections of superior site-specific provenances and
progenies in addition to the establishment of areas of improved
seed production played a critical role. Concurrently, in 1979, the
first clonal plantations were established in the State of Espirito
Santo(Ferreira,1992).Alltheseadvancesledtheareaofeucalypt
plantations to surpass that of pine plantations (Fig. 1a) by the
endofthe1970s.
Thefollowingperiod,1980–2000,wasmarkedbytheconsolida-
tionoftheBrazilianforestsector,involvingmainlybreedingpro-
grams, productivity increases, expansion of cropped areas,
diversificationoftheuseofproducts,increaseofcompetitiveness
and concerns with social and environmental issues (ABRAF,
Fig.1. (a)Annualplantedareawithpineandeucalypt,andtheGrossDomestic
Productsince1961;(b)meanannualincrement(roundwoodwithbark)ofeucalypt 2012). New eucalypt populations emerged from interspecific
andpineplantationssince1990.Plantedareaswascompiledfromdatapublished crosses,especiallybetweenE.grandisandE.urophyllaseekingseg-
by Andrade (1923, 1928, 1936), Reis (1948), Bastos (1961), Ferreira (1989), regationofindividualswithhighproductivityanddesirablewood
BRACELPA(2004)andinformationobtainedfromBrazilianAssociationofForest
quality(Lemos,2012).Superiorindividualsplantedoncommercial
Plantation Producers (ABRAF). GPD data are from World Bank (http://data-
scale in various regions were selected by cloning techniques. In
bank.worldbank.org/data/home.aspx).Themeanweightedproductivitywascom-
putedasafunctionofplantedarea(ABRAF,2012). addition to forest improvement, there was significant advance of
Pleasecitethisarticleinpressas:Gonçalves,J.L.M.,etal.IntegratinggeneticandsilviculturalstrategiestominimizeabioticandbioticconstraintsinBra-
zilianeucalyptplantations.ForestEcol.Manage.(2013),http://dx.doi.org/10.1016/j.foreco.2012.12.030
J.L.M.Gonçalvesetal./ForestEcologyandManagementxxx(2013)xxx–xxx 3
silvicultural practices, especially regarding production of high- followed by the industrial firewood (26.3%), the wood industry
quality seedlings, use of minimum tillage, integrated control of (18.8%),thecharcoalsector(10%),reconstitutedpanels(7.4%),trea-
weeds,pestsanddiseases,judiciousfertilizationrecommendation, ted wood (0.9%) and others (0.6%). Except for firewood, charcoal
andeffectivecontrolofforestfires(Gonçalvesetal.,2008).Thus,at andindustrializedwoodenpanels,whichoperateinthedomestic
the beginning of the 21st century, mean annual increments ex- market, other products are preferably aimed at foreign markets.
ceeded30m3ha(cid:2)1year(cid:2)1roundwood(Fig.1b). Asignificantvolumeofsecondaryproducts(furniture,paper,floor-
Between the 1980s and late 1990s, there was a decrease fol- ing,frames,ironandsteel,etc.)isalsoexported,underscoringthus
lowedbythestabilizationoftheannualplantedareaofeucalypts the importance of the international market for the forest sector.
(Fig.1a),mainlyduetotheendoftaxincentivesin1986.Suchwas The key trends for the forest sector in the short and mid terms
themarkofanewphaseinthehistoryofintensiveforestryinBra- are: (i) increase forest plantation area; (ii) consolidation of ‘‘new
zil(Moratori,2008).Forestcompaniesallocatedlargeinvestments forest frontiers’’; (iii) partial replacement of pine plantations for
for the maintenance of stands, invested in new plantations and eucalypt in the southern and southeastern regions; (iv) develop-
consolidatedassociationsandcooperativeswithpublicuniversities mentofnewmarkets(productsandbusinesses,suchasbio-refin-
forthedevelopmentofscienceandtechnology(Lemos,2012). eriesandbio-energy)(ABRAF,2012).
ThecurrentstatusoftheBrazilianforestsector,sincetheyear
2000, is characterized by the consolidation of Brazil as a major
international player of the planted forest sector. This phase has 2.Locationofplantations
beencharacterizedbytheexpansionofplantedareasandthecon-
solidation of the technological development of the sector, which In2011,Brazil’s totalarea offorest plantationsgrowingeuca-
has been accompanied by a rapid GDP growth in Brazil (Fig. 1a). lyptandpinecovered6.5Mha,being74.8%plantedwitheucalypt
The importance for society of eucalypt plantations and related (Fig.2)and25.2%withpine.Partofthepineplantationsarebeing
industrycanbemeasuredbyassessingsomemainindicators,for replaced by eucalypt. During 2005–2011, the area planted with
instance,roundwoodproductionandconsumption.Itisestimated pine reduced by 189,593ha ((cid:2)1.8% p.a.). The area cropped with
that the potential roundwood production of pine, eucalypt and otherforestspeciessuchasAcaciamearnsii,Acaciamangium,Hevea
teakamountstoaround255.4millionm3yr(cid:2)1.Fromthisamount, brasiliensis (Rubber tree), Schizolobium amazonicum (Parica) and
76.5% corresponds to eucalypt, 23.1% pine, and teak accounts for Tectona grandis (Teak), account for more than 421,588ha, repre-
only0.4%ofthetotalproduction.Themajorpartofthepotential senting6.0%ofthetotalareaofplantedforests(ABRAF,2012).
eucalypt production is concentrated in the Southeast (59.3%), in Thelargesteucalyptplantationsarefoundinthesoutheastern
viewofthesignificantnumberofpulpandpapermillsandmetal- region(56%),especiallyinthestatesofMinasGeraisandSãoPaulo
lurgy charcoal companiesin this region. In 2011,the roundwood (Table1).Largeplantationscanalsobefoundinthenortheastern
consumptionofforestplantationstotaled170.1millionm3,80.6% region(17%)inthestateofBahia;inthesouthernregion,mainly
corresponded to eucalypt plantation productions. The pulp and inthestateofParana,inthecentral-westernregion(11%),mainly
paper sector stood out as the main consumer (36.1% of total), inthestateofMatoGrossodoSul,andinthenorthernregion(5%)
Fig.2. DistributionofeucalyptplantationsanddifferentnaturalbiomesinBrazil(modifiedofABRAF(2012)).FordescriptionofclimatetypesseeTable2.
Pleasecitethisarticleinpressas:Gonçalves,J.L.M.,etal.IntegratinggeneticandsilviculturalstrategiestominimizeabioticandbioticconstraintsinBra-
zilianeucalyptplantations.ForestEcol.Manage.(2013),http://dx.doi.org/10.1016/j.foreco.2012.12.030
4 J.L.M.Gonçalvesetal./ForestEcologyandManagementxxx(2013)xxx–xxx
Table1
Plantedareaofeucalyptin2011,orderandsuborder,texture,localtopographyandextensionofthesoilsindifferentBrazilianregions.
Region Plantedarea Soil
1000ha %Total Orderandsuborder Texture Topographya Plantedarea
1000ha %Region
Southeast 2640 54.2 Oxisols Sandclayloamtoclayey Flattoundulating 1499 56.8
Ultisols Sandclayloam(A-hor)/sand Undulatingtohilly 598 22.7
claytoclay(B-hor)
Inceptisols Clayloamtoclay Rollingtohilly 387 14.7
Entisols
Orthents Clayloamtoclay Hilly 118 4.5
Psamments Sandtoloamsand Flattogentlysloping 38 1.4
Northeast 800 16.4 Oxisols Sandclayloamtoclay Flattoundulating 254 31.7
Ultisols Loamsandtosandclay Undulatingtorolling 400 50.1
loam(A-hor)/sandclay
loamtoclay(B-hor)
Entisols
Psamments Sandtoloamsand Flattogentlysloping 64 8.0
Inceptisols Clay Hilly 54 6.8
Oxisols(plinthic) Clayloam Flattogentlysloping 27 3.4
South 573 11.8 Inceptisols Clay Hillytosteep 225 39.3
Ultisols Sandclayloam(A-hor)/sand Undulatingtohilly 131 22.9
clay(B-hor)
Oxisols Sandclayloamtoclay Undulating 128 22.3
Entisols
Orthents Clayloam Hillytoverysteep 54 9.3
Psamments Sand Flattogentlysloping 6 1.0
Alfisols Sandclayloam(A-hor)/clay Flattoundulating 29 5.1
(B-hor)
Midwest 659 13.5 Oxisols Sandclaytoclay Flattoundulating 313 47.5
Entisols
Psamments Sand Flattogentlysloping 289 43.7
Orthents Clay Hilly 5 0.7
Ultisols Sandyclayloam(A-hor)/ Undulatingtorolling 34 5.2
clayey(B-hor)
Inceptisols Clayey Hillytosteep 13 1.9
Alfisols Loamsand(A-hor)/clay(B- Flattogentlysloping 6 0.9
hor)
North 201 4.1 Oxisols Sandclayloamtoclay Flattoundulating 127 63.0
Ultisols Loamsand(A-hor)/sand Undulatingtorolling 43 21.5
clayloam(B-hor)
Entisols
Psamments Sand Flattogentlysloping 19 9.4
Orthents Clayloam Hillytoverysteep 9 4.3
Oxisols(plinthic) Sandclayloam Undulating 4 1.8
a Topography:0–2%flat,2–5%gentlysloping,5–8%undulating,8–16%rolling,16–30%hilly,30–45%steep,and>45%verysteep(SoilSurveyStaff,1993).
inthestateofAmapa(ABRAF,2012).Thelargestconcentrationof ers,theindustryaimstoensuretimbersupply,reducetheamount
forestplantationsoccursinthesouthandsoutheastofBrazilwhere ofequitycapitalinfixedassets,promotesupportiveprogramsfor
the main industries of pulp, paper, wood panels and steel are employment generation and increase the local income (ABRAF,
located.However,duetocurrentandhighlandpricesinconsoli- 2012).
dated markets of the southern (Parana and Santa Catarina) and
southeast regions (São Paulo), Brazilian forestry is advancing to 3.Edaphoclimaticandphysiographiccharacterizations
otherregionscalledthe‘newforestfrontiers’,asithasalreadybeen
observed in the last 3years in the states of Mato Grosso do Sul, 3.1.Climate
Maranhão,PiauíandTocantins.
Closely related to rainfall and temperature conditions and cli- TheclimaticclassificationofKöppen(1936)hasbeeneffective
mate seasonality, the major biomes where eucalypt plantations indistinguishingthedifferentclimatictypesandsubtypesthatoc-
wereestablishedaretheAtlanticForest(60%),theCerrado(Brazil- curinBrazil,whichhaveclosecorrelationwiththerecommenda-
ian Savannah) with 21%, the Amazon Rainforest (10%) and the tions of eucalypt genotypes and forest productivity. According to
Pampa (lowland grasslands) with 7% (Fig. 2). Caatinga biome thisclassification,wherearetheeucalyptplantations,twoclimate
(Steppe-likesavannah)comprisesonly1%ofeucalyptplantations. typesoccur,tropicalandsubtropical,whicharesubdividedintose-
Previously,grasslandandagriculturalcropsoccupiedmostofthe ven subtypes: fully humid tropical (Af), monsoon (Am), humid
area. tropical with dry winter (Aw), fully humid subtropical with hot
The eucalypt plantations established in lands belonging to summer (Cfa), fully humid subtropical with temperate summer
industrialcompanieshavebeendecreasingoverthelastfewyears, (Cfb),humidsubtropicalwithdrywinterandhotsummer(Cwa),
and in 2010, they represented approximately 73%. Plantations in humid subtropical with dry winter and temperate summer
leased and fostered areas doubled in the last 5years. In 2010, (Cwb). In the Amazon Forest, Af and Am subtypes predominate,
12%ofplantationswereestablishedonleasedlandsand15%infor- in the Cerrado,Aw and Cwa subtypes, and in the Atlantic Forest,
estoutgrowerscheme.Inthismodelofrelationshipwithlandown- Cfa,Cfb,CwaandCwbsubtypes(Fig.2andTable2).
Pleasecitethisarticleinpressas:Gonçalves,J.L.M.,etal.IntegratinggeneticandsilviculturalstrategiestominimizeabioticandbioticconstraintsinBra-
zilianeucalyptplantations.ForestEcol.Manage.(2013),http://dx.doi.org/10.1016/j.foreco.2012.12.030
J.L.M.Gonçalvesetal./ForestEcologyandManagementxxx(2013)xxx–xxx 5
Environmental stresses notably hamper productivity. In cli- weatheringarelow.Theparticlesizedistributionisfinerthansan-
mates with marked seasonality, such as the climatic subtypes dyloam(loamytoclayeytexture).Usually,thesoilmasshasapor-
Aw, Cfb, Cwa and Cwb, productivity is highly variable. Effects of ous massive aspect with strong aggregation of particles into
waterstressare moretypical inAw and Cwa,and thermalstress granules, occurring, less frequently, subangular blocky soil-struc-
(mainlyfrost)inCfbandCwb.WaterstressisnotobservedinAf ture.IntheSouthofSaoPaulostate,Alvaresetal.(2011a)studied
and Cfb. In Aw and Cwa, due to rainfall badly distributed among soilsofwideareasofeucalyptplantationsandfoundhighvariabil-
theseasons,waterstressismediumtohigh.InCfbandCwb,aver- ityofphysicalandchemicalcharacteristicsofOxisolsandrecom-
agetemperatures,andthusevapotranspiration,arelower,because mended specific management of the soil. Small areas (0.7%) of
ofhighlands(Table2).Themostseverewaterstressoccursduring Plinthic Oxisols with sandy clay loam texture and undulating
periods of Indian summers, when there are 2 or more weeks of topographyareplantedinthenorth.
drought in the middle of the rainy season, especially if average Ultisolsalsoappearasanimportantsoilusedforeucalyptplan-
temperaturesarehigh. tations, with 1.2Mha (25.5%), mainly southeastern, northeastern
The first eucalypt plantations were established in temperate and southern Brazil. They are deep to shallow, well to poorly
zonesofSoutheasternandSouthernstates.Inrecentdecades,the drained,withsignificantilluviationofclayfromthesurfacehori-
plantationshaveexpandedtomoretropicalregionsoftheNorth- zonstotheBhorizon,evidencedbythetexturalgradient(erosion
eastandCentral-West,especiallyinAmandAwclimates.Inthese susceptibilityincreaseswiththetexturaldifferences).Specialcare
regionsofinexpensivelandbutwithadryseasonwithhighwater relatedtosoilmanagementshouldbetakenwhenAhorizonissan-
stress,themajorchallengeistoobtainnewgenotypesadaptedto dyandBhorizonisclayey,whichcommonlyoccursinareaswith
thesewarmerandxericconditions. steep slope. The B horizon usually has faces of peds with shiny
waxysheen,andithasmediumfertility.
3.2.Soilandtopography Approximately 0.68Mha (14%) of eucalypt plantations are
growninInceptisols,mainlyinthesoutheastandsouth.Theyare
An estimate of the major Brazilian soil and topography types moderate to well drained, shallow, rarely deep, with pedogenic
usedforeucalyptplantationswasperformedusingseveralsources development littlepronounced.The contents of easily weathered
of spatial and tabular data (Table 1). Data for each area was primarymineralsaregreaterthan4%,beingcommonthepresence
adoptedfromABRAF(2012).Theareamappingsofplantationsin ofrockfragments.Theirtexturerangesfromclayeyloamtoclayey.
the states of Sao Paulo (IF, 2002) and Minas Gerais (Scolforo Physicalconstraints(permeability,depth,waterretention)limiting
et al., 2008) were arranged in a single database. These states are eucalyptproductionaregenerallyfound,withoutnecessarilybeing
the largest eucalypt planters, accounting altogether for 51.5% of economicallyunfeasible.Commonly,Inceptisolsareofmediumto
thetotalplantedarea.Theremainingarea(fourteenotherstates) highfertility.
was joined in tabular information for municipality-based forest Thelittledevelopedsoils,suchasLithicOrthentandEntisolsde-
plantations (IBGE, 2011). Orders and suborders of soils were ob- veloponsands(Psamments)presentAhorizonoverlyingrockorC
tainedfromthepedologicalmapofBrazil(IBGE,2001).Suchmap horizon.Orthenthaslowdepth,buthighclaycontentandgener-
is classified according to the Brazilian soil taxonomy (EMBRAPA, allymediumtohighfertility.Thereareapproximately185,000ha
2006),whichwasconvertedtotheUSsoiltaxonomy(SoilSurvey (3.9%)cultivated mainly in the southeastern and southernstates.
Staff, 1993). The map of clay content produced by Silva et al. Thetopographyisstronglytilted,rangingfromhillytoverysteep.
(2011) was used as information source of clay content. The map MostlandscoveredbyOrthenthavephysicalcharacteristicsunfa-
of clay content was interpolated by the kriging method of 5500 vorabletoeucalyptdevelopment andare locatedin areasofper-
Braziliansoilprofiles.Usingageographicinformationsystem,Alv- manent preservation of natural vegetation. Psamments are
aresetal.(2011b)compiledadigitalelevationmodelfromShuttle mostly cultivated in the central-western region, in flat to gently
RadarTopographyMission(SRTM)(FarrandKobrick,2000)inits sloping topography. In the State of Rio Grande do Sul, there are
current fourth version (Jarvis et al., 2008). Simple geoprocessing extensive areas of Alfisols cultivated in flat topography and high
techniques(Ormsbyetal.,2010)wereusedtoperformintercepts texture relation. These soils usually have sandy clay loam (A-
of clay contents between the map of eucalypt plantations and hor)/clayey (B-hor) texture. In many places, plantations show ef-
thepedologicalmap,andtheslopemaptoelaborateTable1. fectsofanoxiainthesesoils.
Thewidevariabilityofsoilsandtopographyreflectsthebroad
rangeoftheplantingareaaswellastheactionandthedominance
ofvariousfactorsandprocessesforsoilformation(Tables1and2).
Generally, the most weathered and developed soils (oxic B hori- 4.Geneticadaptation
zon) are under very active climatic and biotic conditions, as in
the case of tropical and subtropical humid lands that are located ThreephasescharacterizetheeucalyptscultureinBrazil,under
inthelandscapeatmorestablepositions,flattoundulatingtopog- treebreedingperspectives.
raphy.Moderatelydeveloped,lessweatheredsoils(argilicBhori- Thefirstphasewasfromeucalyptintroductionatthebeginning
zon)areformedatlessstabletopography(undulatingtorolling). oflastcenturybyNavarrodeAndradetosupplyfirewoodpolesand
Soils less developed than the previous ones (cambic B horizon) railwaysleepersuntiltheadventoffiscalincentivesforreforesta-
are formed in rolling tovery steep topography. Poorlydeveloped tioninthelate1960s.Atthisstage,thelarge-scaleeucalyptplan-
soilsarecommonlyfoundinveryunstabletopography. tationswererestrictedtoSaoPaulostate.ThemainspecieswereE.
Oxisolisthemainsoilorderoccupyingalmosthalfoftheland grandis,Corymbiacitriodora,Eucalyptuscamaldulensis,E.saligna,E.
used for eucalypt plantations (47.9%). This soil type is found in urophylla (known then as E. alba). The second phase, with fiscal
allregions,butespeciallyintheSoutheast,inSãoPauloandMinas incentives, lasted until the 1980s and was characterized by the
Geraisstates.Itisdeep,withwell-differentiatedhorizonsandlow expansion of planted areas using species such as E. grandis. The
texture gradient. Due to the advanced stage of weathering and useofseedsfromselectedorigins,suchasCoffsHarbour,Australia,
intensive leaching process, the predominant clays are kaolinitic generated in seed production areas (SPA’s) brought significant
andoxidic(gibbsite,goethiteandhematite).Thecationexchange gainsinproductivityinthestateofSaoPaulo.However,eucalypt
capacityislowerthan13cmol kg(cid:2)1clay(aftercorrectionforcar- plantations established in other regions using seeds selected for
c
bon), fertility and content of minerals with low resistance to theSaoPaulostateshowedproblemsofadaptationtobioticfactors
Pleasecitethisarticleinpressas:Gonçalves,J.L.M.,etal.IntegratinggeneticandsilviculturalstrategiestominimizeabioticandbioticconstraintsinBra-
zilianeucalyptplantations.ForestEcol.Manage.(2013),http://dx.doi.org/10.1016/j.foreco.2012.12.030
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nage.(2013),http://dx.doi.org/10s,J.L.M.,etal.Integratinggenetic OMDNRMecraeeygcttw(aaiiuemvnonsirmeentrmaahaepvnnsnelyeonnecrgensuueaaesaat(rtoull5na(cid:2)frtr0o1eifa).ormi(omnn(cid:3)smfCota)nlrltahinsfall) 22NNAN04toool0–nnrat02neeh–6tei3ca0sF0to0(rceosats(tsaml)allarea) 12SNANh54mooo0–nrart02ehtz–6o(2(n6e0a30Fs)o0treersntpart) M2SFNNCM1oa10eooeivrr0–dnrderat02weasihn–5utdeen2moasa0tsh(0ttB()o0m(rawahnoziesdigslthiltAayent()rl3naa–)nn7td)ic Aa1SLS(1sinho65tgoludo–0ahurt02nttCht–0htea(i2eacr6n5rnrdnF2a0dod)0fmrrSoeeoosqustutt(h)emneatossttly) NA1S(S1hoe35toilvu–0gnaet01hnehr–7lteai2acna5ndnF0dsod0rSofeorfsuetcqtohuaeesants)tt M2ASL1io00tgelu–0ahdt02ntih–2utaei1mnca5sdF0t(oi60nr3efrs–et5q)uent iSh1ASM1nho80itfgoluo0–rahdert02nlhteq–0atreu(in1ac6ae8dtsneF30stot))0a(rinendstterior J.L.M.Gonçalvesetal./ForestEcology
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in mediumdrainage,acidic,poor mediumdrainage, aerated,mediumand aerated,mediumtoslow mediumdrainage, mediumtorapid mediumtorapid x
im supplyofplantnutrients(SPN) acidic,poorSPN rapiddrainage,acidic, drainage,acidic,medium acidic,mediumto drainage,acidic, drainage,acidic,
iz poorSPN SPN excellentSPN poorSPN mediumSPN
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ab a TypesandsubtypesaccordingtotheclimaticclassificationofKöppen(1936).
iotic b Topography:0–2%flat,2–5%gentlysloping,5–8%undulating,8–16%rolling,16–30%hilly,30–45%steep,and>45%verysteep.
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J.L.M.Gonçalvesetal./ForestEcologyandManagementxxx(2013)xxx–xxx 7
suchastrunkcankercausedbythefungusCryphonectriacubensis to produce superior individuals with higher growth, adaptation
andabioticfactors,suchasdroughtperiods. andwoodquality(AssisandMafia,2007).Theuseofclones,rather
Field observations and research showed that these problems thanseedlingsoriginatedfromseeds,allowsthetransferoftheto-
couldbelargelysolvedusingclonesofinterspecifichybrids,such talgeneticvariance,whichincludestheadditivevarianceandthe
asE.urophylla(cid:3)grandis(‘‘urograndis’’),leadingtothecurrentthird varianceduetothedominanceeffects(Zobeletal.,1982).Thispro-
phase. The first hybrids were selected in plantations originated cedureresultsinmaximumgeneticgainsregardingwoodproduc-
from seeds of natural hybrids, spontaneously generated by con- tivity,woodqualitypropertiesandresistancetobioticandabiotic
taminationofseedcollectionareas.TheSimpleRecurrentSelection factors.Clonesfromhybridsaremoreadvantageousthanseedlings
(SRS) was used thereafter as a breeding strategy based on con- ofsinglespecies,becausetheycancombinesilviculturaltraitssuch
trolledpollination.Later,aftertheoccurrenceofheterosisininter- aswoodqualityandadaptability(Table3).
specific crosses be showed (Coterill, 1997; Resende and Resende, AccordingtoAssisandMafia(2007),themostimportanteuca-
2000), the Reciprocal Recurrent Selection (RRS) became one of lyptspeciestogeneratehybridsresistanttodroughtarefoundin
the largely used strategy to generate hybrids. Several breeding theExertariasectionfromnorthernQueensland,Australia(E.cam-
strategies may be used, however the RRS Selection of parents aldulensis, Eucalyptus tereticornis and Eucalyptus brassiana). For
and Generation of Intermediate Hybrid (RRS-G-IH), proposed by siteswherefrostsarefrequent,therecommendedspeciesforhy-
Resende and Higa (1990), is one of the most suitable because it brid generation are: Eucalyptus viminalis, Eucalyptus badjensis,
leadstogreatergeneticgainpertimeunit,andsuperiorintermedi- Eucalyptus benthamii, Eucalyptus smithii and Eucalyptus dunnii. In
ate hybrids before the RRS cycle is completed. The RRS-G-IH is practice, subtropical species of eucalypts exhibit low rooting
basedontheselectionofparentsofeitherspeciesorpopulations, capacity,makingitdifficulttousethemaspurespecies.Therefore,
on hybrid progeny tests and on recommendations of the parents itisrecommendedtocrossthemwithspeciesthathavegoodroot-
themselves (a pollen mix) to generate a new hybrid population. ingcapacity.
The intermediate hybrids (IH) are obtained by crossing parents The hybrid E. grandis(cid:3)urophylla is not fully adapted to very
with high General Ability for Hybridization (GAH) that had not warm and humid climates as in northernBrazil(Amazonregion)
been previously crossed, during the generation of hybrid proge- in particular due to leaf diseases (Hardiyanto and Tridasa, 2000).
nies.Thisnewpopulationofhybridsisobtainedconcurrentlywith Creatinghighlyproductiveeucalyptmaterialremainsachallenge
the recombination of the parents; therefore it is a process of in northern Brazil where growth rates are still far lower than in
obtainingnewhybridsbeforeanewRRScycle(Fig.3). south-eastern Brazil (Souza et al., 2004; Demolinari et al., 2007;
Theinterspecifichybridizationisaquickandefficientmethod Behlingetal.,2011).Eucalyptuspellita(F.Muell.)isoneofthemost
to obtain gains in genetic improvementof eucalypt species, with promisingspeciesforeucalyptbreedingprogramsinhumidtropics
significantanddirectimpactonforest-basedindustry.Thesearch (Harwoodetal.,1997;Brawneretal.,2010).Thisspeciesisnative
ofcomplementarytechnologicaltraitsinthewoodcharacteristics, tonorthernQueensland(Australia)andsouthernNewGuineaand
namelytolerancetobioticandabioticstressandmanifestationof isresistanttotomanytropicalpestsanddiseases(Guimarãesetal.,
theheterozygosisasobservedinseveralhybrids,isthemainway 2010).ItislargelyusedinIndonesiaforpulpwoodproduction(Lek-
sono et al., 2006; Brawner et al., 2010) and could be used for
hybridizationwithE.urophylla(VigneronandBouvet,2000).
In addition to improvements of plantation productivity and
woodquality,rootingability,resistancetodrought,frostanddis-
eases, research is also being conducted to increase orthotropism
(stem straightness) and resistance to wind. There are some risks
associatedtoclonalforestry,mostlyrelatedtotherestrictedgenet-
icbaseofplantations.Mostcommercialclonalforestsdonotfollow
recommendationfromtheoreticalstudies,whichhaveshownthat
clonalplantations shoulddeployatleast40 differentclones(Bri-
shirandRoberds,1999).Anotheraspectistheneedofcontinuous
investmentinbreedingprogramstogeneratenewclones.Ingen-
eral,anewclonetakesatleast12–16yearstobegeneratedandap-
provedforcommercialuse.Itisrecommendedthat,onaverage,a
Fig.3. SimplifiedschemeofRRS-G-IH:ReciprocalRecurrentSelection(RRS)with new clone should be introduced every year. Currently in Brazil,
recombination of the parents themselves and use of Intermediate Hybrid (IH)
mostofthebreedingprogramstogeneratenewclonesaredevel-
betweentwoselectionscycles.Where:P1andP2=parentsfrompopulations1and
2;H1andH2=newhybridpopulations;IH=intermediatehybrids;R=recombined oped by pulp and charcoal private companies. Breeding for solid
parentsgeneratedfromcrossingwithineachpopulation. woodproductsislessdeveloped.Theuseoftransgenicgenotypes
Table3
RelativequalityofEucalyptusspeciestraitswhichmaybecombinedintheinterspecifichybrids:VG=verygood;G=good;R=regular;I=intermediate;?=withoutinformation.
Source:Santosetal.(2008).
Species Growth Rustresistance Rooting Canker Droughtresistance Frostresistance Wooddensity Screenedyield Lignin Hemicellulose
E.grandis VG I G I R R R I I I
E.urophylla G I VG VG I R I I R I
E.camaldulensis R VG VG VG VG R VG R R R
E.tereticornis R VG G VG VG R VG R R R
E.robusta I VG VG VG I R VG R R R
E.resinifera I VG VG VG I R VG R R R
E.pellita I VG VG VG I R VG R R R
E.dunnii I I R R R G G G G R
E.benthamii VG G R ? R VG R R R I
E.globulus R I R ? I G VG VG VG VG
Pleasecitethisarticleinpressas:Gonçalves,J.L.M.,etal.IntegratinggeneticandsilviculturalstrategiestominimizeabioticandbioticconstraintsinBra-
zilianeucalyptplantations.ForestEcol.Manage.(2013),http://dx.doi.org/10.1016/j.foreco.2012.12.030
8 J.L.M.Gonçalvesetal./ForestEcologyandManagementxxx(2013)xxx–xxx
isalsobeingstudiedinBrazil.Herbicideresistanceandreduction Thornthwaite andMather,1955), itisrecommendedtoplantthe
ormodificationoflignincontentandcompositionmaybeachieved hybrid E. urophylla(cid:3)grandis, E. grandis and E. urophylla. Among
throughtransgenesis.However,theavailabilityofgeneticmaterial thesethreespecies,E.grandisisthemostsensitivetowaterstress,
resistanttopestsanddiseasesoradaptedtofrostanddroughtcon- andshouldbeplantedindeeperandclayeysoils.E.duniipresents
ditionsinBrazilwilltakeawhiletooccur. problems of seed propagation (late flowering) and cloning (poor
rooting).
Still under subtropical conditions, if there is moderate water
5.Matchingspecies/hybridstosites stress (100–200mmyear(cid:2)1), which occurs in regions with hot
summers,thehybrids,inorderofincreasingdroughttolerance,E.
Thewiderangeofeucalyptspeciesandhybridswithdifferent urophylla(cid:3)grandis, E. grandis(cid:3)camaldulensis (grancam), E. uro-
climaticandedaphicaptitudesassociatedwiththeeasypropaga- phylla(cid:3)camaldulensis(urocam),andthepurespeciesE.urophylla,
tion by seeds and cloning allow the adaptation of plantations to E. camaldulensis and E. tereticornis are most suitable. The hybrids
various tropical and subtropical regions in Brazil. The possibility propagatedbycloningandmorespecificallyrelatedtoedaphocli-
of using eucalypt wood for a range of purposes has stimulated maticadaptation,arethemostplanted.Thisflexibilityofoptions
the establishment of forests for multiple uses by both large and allowsobtainingyieldssimilartothoseinlowwaterstresscondi-
smallenterprises(Table4).Thedesirablecharacteristicsinassoci- tions (Table 5). The E. urophylla(cid:3)grandis is the genetic material
ation with the accumulated knowledge on eucalypt silviculture mostplantedinBrazil,sinceitsedaphoclimaticadaptationiscom-
encouragetheuseofthisgenusinmostplantationsevenmore. patiblewiththeregionswherethelargestplantationareasarelo-
Themostimportantselectivefactorsforagenotypeallocation cated.E.urophyllahashighadaptabilitytoseveralregionsandhas
arewoodcharacteristics,productivitylevel,susceptibilitytopests beenwidelyhybridizedwithE.grandis,aimingtoobtainmaterials
anddiseases,droughttolerance,especiallyintropicalregions(frost toleranttodroughtsandresistanttoeucalyptcanker.AlthoughE.
free), and frost tolerance in subtropical regions (mostly without urophylla and E. camaldulensis present some favorable traits in
waterdeficit). common, E. camaldulensis has greater resistance to drought and
Undertropicalconditions,inregionswithpronouncedseason- higher wood density than E. urophylla as well as more alleles of
alityandmoderatetolongdroughtperiods,theplantingofhybrid importance for hybridization. In regions where E. urophylla is
genotypespredominates,propagatedbycloning.Undersubtropical adapted,it has a higher volumetric growth than E. camaldulensis.
conditions,theplantingofsinglegenotypespredominates,propa- Generally E. urophylla adapts to a wider range of environments
gated by seed (Tables 4 and 5). Clonal plantations withinterspe- than E. camaldulensis (Tables 4 and 5). Both species are good
cifichybridshavebeenfundamentaltoadapteucalyptinregions sourcesofrustresistance,althoughtheintraspecificvariabilityre-
withwaterandnutritionalstresses.Clonesfromhybridsaremore quirespreliminaryassessmentforresistancebymeansofinocula-
advantageousinrelationtoseed-originatedseedlingsofpurespe- tionsundercontrolledconditions(Fonsecaetal.,2010).
cies, because they can combine silvicultural traits such as wood Whenthereishighwaterstress(200–400mmyear(cid:2)1),typical
quality and site adaptability (Tables 3–5). Therefore, cloning is intropicalconditions,Awclimatetype,withwarmanddrywinter,
essential for the fixationof desirable genotypecombinations and thehybridsarethemostrecommendedgenotypes.AmongthemE.
hybrid vigor as it allows obtaining homogeneous forest products grandis(cid:3)camaldulensis, E. urophylla(cid:3)camaldulensis, E. tereticor-
andmaximizationofselection(Fonsecaetal.,2010). nis(cid:3)brassiana and E. urophylla(cid:3)tereticornis are highlighted. It is
Genotypes propagated by cloning, due to the small genetic alsorecommendedtheuseofpurespeciesE.camaldulensis,E.teret-
diversity,arelittleplasticrelativelygenotypespropagatedthrough icornisandE.brassiana(Tables2and5).Mostareaswiththesecli-
seeds. Hence, they have a higher risk of genotype-environment mate conditions are located in the central-western and
incompatibility (Gonçalveset al., 2011). Thus,the specific alloca- northeastern regions (western), where the ‘‘new forest frontiers’’
tion of a genotype to a site should be well tested, based on field arefound.Thereisnotmuchknowledgeofabioticandbioticinter-
andlaboratorytests.Theuseofclonesinforest-basedenterprises actionsofgenotypeswiththesite.Above400mmyear(cid:2)1ofwater
istheresultofextensiveinvestmentsinresearchandpilotplanta- deficiteucalyptplantationsarenoteconomicallyviable.
tionsinregionswithdifferentedaphoclimaticcharacteristics(Higa GavaandGonçalves(2008),Alvares(2011)andGonçalvesetal.
and Silva, 2008). Given the rapid advances in eucalypt breeding, (2012)evaluatedtheeffectofdifferentphysicalandchemicalsoil
regarding adaptation to water stress and resistance to pests and attributes on wood quality and productivity of E. grandis and E.
diseases,andtheadoptionofclonalpropagationtechniques,geno- grandis(cid:3)urophylla plantations for cellulose productionin several
typesrapidlybecomeobsoleteandarereplacedbymoreproduc- sites at the Western Plateau of the State of São Paulo, with ages
tiveonesafterharvesting.Thus,thereplantingofcropsbecamea ranging between 6.0 and 7.0years. Four soil types, with texture
common procedure after the second half of the 1990s in Brazil, ranging from sandy to very clayey were found. They found high
whilecoppicewasmorefrequentbefore(Fonsecaetal.,2010). relationship between MAI and several attributes of soil, mainly
Undersubtropicalconditionsinregionswithhotsummerand clay content, which was directly related to the amount of soil
good rainfall distribution, Cfa climate type, with no more than a plant-available water content (Fig. 4; see also Marsden et al.,
monthofwaterdeficit,andlightandfrequentfrost,themostrec- 2012,thisissue).GavaandGonçalves(2008)foundthatbasicwood
ommendedgeneticmaterialsarethehybridE.urophylla(cid:3)grandis densitydidnotchangedatdifferentsoiltypes,totallignincontent
(urograndis), E. grandis, E. urophylla, E. saligna and C. citriodora. If decreased and holocellulose content and screened cellulose yield
thesummeristemperate,Cfbclimatetype,whichoccursinsouth- increasedexponentiallyassoilclaycontentincreased(untilabout
ern Braziland high-altitudeareas ofthe southeast,thespecies E. 350–400gkg(cid:2)1ofclay).Thewoodextractivescontentwasnotaf-
dunii,E.benthamiiandE.salignaareused(Tables2and5).E.dunii fectedbysoilattributes.
toleratesmoderatefrostandE.salignatoleratemildfrost,sothey
should be planted in the higher quotas of the terrain. In areas of
lowerquotas,middlethirdoftheslopeandlowlands,wherefrost 6.Rootconfigurationdependingonwaterandnutrientstresses
is severe and frequent, it is recommended to use E. benthamii. If
the summer is mild and the winter is dry, Cwb and Am climate Traits of the eucalypt and its outstanding success to thrive in
types, with low water stress (i.e., a water deficit of 50– harsh environments reflect the evolution of a particularly strong
100mmyear(cid:2)1, according to the soil water balance proposed by adaptation where water and nutrients are limited. There are a
Pleasecitethisarticleinpressas:Gonçalves,J.L.M.,etal.IntegratinggeneticandsilviculturalstrategiestominimizeabioticandbioticconstraintsinBra-
zilianeucalyptplantations.ForestEcol.Manage.(2013),http://dx.doi.org/10.1016/j.foreco.2012.12.030
zP
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pa
lantations.Forticleinpress ChaSrpaecctieersis/htiycsbroidfswood,mainuses(W,gsocuomsdc(cid:2)ed3p)etinbsiiltityy/resisLctaoignnnctieenntto(p%e)stsanMdadiinseuasseessandsproutingcapaSPcueitssytcseobpftsibpielcitiyes/raensidsthaynbcreidscommonlyusedinplantatiDonissea(ssheoscrtrotation6–8years). CsSptaurpomaucptiitnygoafVigorof
restEas:G Corymbiacitriodora 0.72–0.75 22–25 Sawnwood,fuelwood, Susceptibletobluegumpsyllid(BGP);tolerantto SusceptibletoCylindrocladiumleafspot; Lsuorwvitvoal Lshoowotto
coon charcoal,treatedwood redgumlerppsyllid(RGLP),bluegumchalcid(BGC) resistanttocankerandveryresistanttorust; medium medium
l.Maçalve E.benthamii 0.48 24–25 Pulpwood,sawnwood, SaunsdcbeprotinbzleebtougBG(BPB) sVuesrcyerpetsibislteantotAtormruilslatraiandrocoatndkiesrease(ARD) Lowto Medium
nages,J.L E.camaldulensis 0.60–0.70 28–32 tPruelaptwedoowdo,osdaw,finrwewooodo,d Susceptibletoseveralspeciesofpsyllid;very SusceptibletoleafspotofMycosphaerella mMeeddiiuumm Medium J.L.M
.(2013),http://dx.doi.org/10.M.,etal.Integratinggenetic EEEE....dgstearuralniengntndiiciiaosrnis 0000....44464260––––0000....65455282 22232440––––32222367 ctswpPPPPrahuuuuaeowanllllaoppppretndcwwwwelwo,dooooacooooolhwoddddado,,,,r,ocsfssupodaaaeawwwa,lnlfiwnnnerwwwleo,wooootdrooooe,dddoa,,,dted sSVtVVouueeessrrrRccyyyGeesssppLuuuPttsissibbcccleleeeeppptttttioiiobbblRllBeeeGGtttLPoooPBa,BRnBBBGd;BLasPBaun,BnsdBcdGeBBpGCGtCiabC;nltedotlBoeBrBaGntC;totoRleGrLaPnt (rsScs(RSGLseuuauueoeSsnsssssoviMcccciksdeseeeetet)rapppprasan(;ttttonlsiiiitusbbtbbesutllvrplltoeeeecseoeecrrttcttesruaooooiupelsoLcssvLttsfiatS)Sepab;nrMMrnelevykecdseei((istercsrsMuoy;setasa).vnsLr;cnemukSeercrMesspeaaeicrstsrl;eiitkt(bsspsoassptulteinaeiriesvtbnuacctelttsinoeereotpadt;srtoltc)uoviMasbvsepnLtle.reSekyraMcyetniored;sA)RD tLmLmMtMooooeeeewwhhddddiiiiiiggttuuuuoohhmmmm tMLmMtMoooeeeewhhddddiiiiiiggtuuuuohhmmmm .Gonçalvesetal./ForestEcology
.1016ands panel mreosillsetrainancea)t;orecasinsktaenrtvtaoriveesrfyrormesivsetarynttorust; tohigh tohigh andM
/j.foreco.20ilvicultural EE..cuarompahlydlulalensis(cid:3)grandis 00..4478––00..4596 2297––3219 tPFruueelapltwwedoooowddo,,ocfuhdea,lrpwcuoolapolwd,,ood VVpBsGeeyrrClyylidssuu(ssCcc.eespppttaiitbbulleleatttaooaRBnGGdLCPB;;.ssuoucssccciedepeptntitbiballeliest)to;otBEoBulecaraanlnydptttuos SsGsuuuosssocccdeeepppstttoiiiubbbllrleeecetttsoooohLcfSaigrMnehksle(yissretravenseicrsaetlatsnoptreucsietsa)ndcanker; HHiigghh HHiigghh anagementx
12.12.030strategiesto EE..uurroopphhyyllllaa(cid:3)(cid:3)ggrloabnudliuss 00..4581––00..5524 2263––2295 fPPfiuuureelllppwwwwoooooooodddd,,,cphcahanarecrloc,oaall, SRUuGnsLicdPeepnttiibfileedtoocRcGuLrrPenancediBnBtheplantations sGRueossocidsetpastoniubtlrtecoetsrouoLsftSrMaens(dissetTavenercaraetoltssopphreaucesiretisaa)nsdppc.anker; tMtMooeehhddiiiigguuhhmm MtMtooeehhddiiiigguuhhmm xx(2013)xxx–
minim E.urophylla(cid:3)tereticornis 0.53–0.64 28–35 sFauwelnwwoooodd,,pturlepawteododw,ood VerysusceptibletoRGLP SusceptibletoleafspotofCylindrocladium High High xxx
iz charcoal pteridis
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ab ThistablewascompiledfromdatapublishedbyHigaandSturion(1991),Ferreira(1992),Pereiraetal.(2000),Fonsecaetal.(2010),andinformationobtainedfromforestcompaniesassociatedtoForestryScienceandResearch
io Institute(IPEF,Brazil).
tic a Stumpsurvival:high(>90%),medium(80–90%),low(<80%),mainlydependentonthegeneticcharacteristicofthespecies/hybridandenvironmentalconditions.Generally,thegreaterthewaterdeficit,theloweristhecapacity
an ofsprouting.Ifsurvivalislow,itisnotrecommendedusecoppicesystem.
d b Pests:bronzebug(Thaumastocorisperegrinus);bluegumchalcid(Leptocybeinvasa);redgumlerppsyllid(Glycaspisbrimblecombei);Eucalyptuspsyllid(CtenarytainaspatulataorBlastopsyllaoccidentalis);bluegumpsyllid
b
io (Ctenarytainaeucalypti).
tic c Diseases:Eucalyptusrust(Pucciniapsidii);basalcanker(Diaporthecubensis);Eucalyptusleafdisease(Mycosphaerella);Armillariarootdisease(Armillarialuteobubalina).
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10 J.L.M.Gonçalvesetal./ForestEcologyandManagementxxx(2013)xxx–xxx
Table5
Climatetype,meanannualrainfall,temperatureandactualevapotranspiration,dryseason,themainspeciesandhybridsrecommendedforplantingandexpectedaverage
productivity(roundwoodwithbark)oftheplantedareaswitheucalypts.a
Typeclimate Meanannual Meanannual Meanactual Dryseason Species/hybridc Meanannualincrement
rainfall temperature evapotranspiration Numberof Waterdeficitb (m3ha(cid:2)1year(cid:2)1)
(mmyear(cid:2)1) ((cid:3)C) (mmyear(cid:2)1) months (mmyear(cid:2)1)
Cfa,Cfb 1500–2500 13–20 500–1000 0–2 0–50 EUG,Egr,Eur,Esa,Cci,Edu,Ebe,EUG 35–60
Cwb,Am 1000–1800 18–20 800–1100 2–3 50–100 EUG,Egr,Eur 35–45
Cwa,Aw 1000–1800 20–24 1000–1200 3–4 100–200 EUG,Eur,EGC,EUC,Eca,Ete 35–45
Aw 1100–2000 24–26 1100–1500 4–6 200–400 EGC,EUC,ETB,Eca,Ete,Ebr,EUT 25–35
As,BSh 700–1500 23–27 600–1000 >6 >400 Plantingisnotfeasible
a ThistablewaspreparedbasedoninformationobtainedfromforesterswhoworkforcompaniesassociatedwiththeInstituteofForestResearchandStudies,Brazil,and
informationpresentedbyFonsecaetal.(2010)andSilva(2008).
b AccordingtothesoilwaterbalanceproposedbyThornthwaiteandMather(1955).
c Egr=E.grandis;Esa=E.saligna;Eur=E.urophylla;Cci=Corymbiacitriodora;Ete=E.tereticornis;Ebr=E.brassiana;Ebe=E.benthamii;Edu=E.dunnii;EGU=Eucalyptus
urophylla(cid:3)grandis(urograndis);EGC=E.grandis(cid:3)camaldulensis(grancam);EUC=E.urophylla(cid:3)camaldulensis(urocam);ETB=E.tereticornis(cid:3)brassiana;EUG=E.urophy-
lla(cid:3)globulus;EUT=E.urophylla(cid:3)tereticornis.
to site. Species or hybrids exhibiting low drought tolerance pro-
ducealargenumberofmediumandfinelateralrootinthesurface
soilhorizons,butnotadeepandstrongtaproot,andtherootsys-
temsshowasmallresponseordonotrespondatalltodrysoils.
Moreover, highly-drought tolerant species produce vigorous and
deep tap roots, and sinkers emerging from the laterals towards
thebaseofthetubeinadrysoil.Theabilityofthegenotypetorap-
idlydevelopastrong,deeplydescendingrootsysteminadrysoilis
undoubtedlyanimportantadaptivefeatureindroughtconditions
(Krejci,1986;GonçalvesandMello,2004;Florence,2007).
IneucalyptplantationsestablishedincoastalregionsofBrazil,
inclayeycohesiveUltisolswithphysicalimpedimentslikefragipan
andiropan,underhighwaterstress,speciesandhybridsmoretol-
erant to droughts are those that produce the more vigorous tap
roots associated with the capacity to explore cracks by sinkers.
Forthegenotypicadaptationof eucalyptin these edaphoclimatic
conditions,thedevelopmentofspecificequipmentandtechniques
specificfordeepsubsoiling(80–120cm)was fundamental(Stape
etal.,2002;Souza,2002).
Fig. 4. Relationship between increases in mean annual increment (MAI) of
Most total root length of eucalypt plantations consist of fine
roundwoodandclaycontent(0–20cmlayer)forthreesitesofeucalyptplantations
inSaoPaulostate. roots,andmainlyestablishedintheuppersectionofthesoilprofile
(GonçalvesandMello,2004).Itisnotonlythedistributionoffine
rootsinthesoilprofilethatcontributestotheadaptationofageno-
numberofattributesoftreerootsthatinfluencetheabilityofthe typetoitsenvironment,butalsotherootdynamics.Finerootpro-
tree to compete for soil resources in regions of small availability ductions exhibit strong seasonal variations (Laclau et al., 2004;
of water, nutrients, air, growing space (Gonçalves and Mello, Mello et al., 2007; Thongo-M’Bou et al., 2008). When edaphocli-
2004;Florence,2007). Theattributesincludethestructureofthe matic conditions are favorable, there is rapid production of fine
root system, the distribution of fine roots, the seasonality of roots,fromrecentassimilatesorfromcarbohydratesandnutrients
growth,and the physiological ability to take up waterand nutri- stored in thicker roots (Ford and Deans, 1977). In times of in-
ents(Comerfordetal.,2006). creasedwateravailabilityandsoiltemperature,theproductionof
Therootstructureandgrowthareimportantattributesgeneti- finerootsisaccelerated,boostingshootgrowth(TeskeyandHinck-
callycontrolledbythespeciesandindividual—anevolutionaryre- ley, 1981; Hendricks et al., 1993; Joslin et al., 2001; Mello et al.,
sponse to maintain the competitive ability in sites with specific 2007). There is continual death and replacement of fine roots
characteristics (Florence, 2007). In general, mechanisms and pro- (Thongo-M’Bou et al., 2008). Observations of the root system at
cessesthatenableaplanttocopewithperiodic,severe,waterdef- anytime provide only a static view of the system. The dynamic
icitinthesoilinvolveatradeofftotheplantintermsofreduced view is much more valuable, particularly when it includes the
growthpotential(GonçalvesandMello,2004). turnover of fine roots during the year, the soil exploration rate
The eucalypt root system has high relationship with the site byroots,andthetimesoftheyearwhenrootsystemsaremostac-
quality. This may be expressed through the root: shoot ratio of tive(Melloetal.,2007;Jourdanetal.,2008;Laclauetal.,2010a,b).
theplant,oratendencytodevelopastrongtaprootaccordingto A more dynamic root system is more effective to explore a large
waterstress.Thetreerootconfigurationmaybemodifiedaccord- soilvolumeandmaximizeaccesstowaterandnutrients.
ingtositeconditions(soilmoisture,depth,textureandstructure, Thesenescenceoffinerootscanbeacceleratedbystressfulcon-
stoniness,hardhorizons,etc.)(Jacobs,1955;Krejci,1986;Gonçal- ditionsduetoseasonalvariationsintemperature,soilwaterstatus,
vesandMello,2004).Thishashighvariabilityintermsofspecies injuries to canopy (defoliation, parasitism), hormonal changes,
and hybrids. Species and hybrids that express different degrees among others. Thus, fine roots are ephemeral components and
of drought tolerance show differences in the root structure. Tap haveaturnoverratecomparabletoleaves;therefore,itisexpected
root depth, vertical and lateral distribution of roots, particularly similar response patterns of these components to environmental
finefeederrootsplayasignificantroleindeterminingadaptation factors (Lyr and Hoffmann, 1967; Gonçalves and Mello, 2004).
Pleasecitethisarticleinpressas:Gonçalves,J.L.M.,etal.IntegratinggeneticandsilviculturalstrategiestominimizeabioticandbioticconstraintsinBra-
zilianeucalyptplantations.ForestEcol.Manage.(2013),http://dx.doi.org/10.1016/j.foreco.2012.12.030