Exotic tree Artocarpus heterophyllus (Moraceae) invades the Brazilian Atlantic Rainforest / Árvore exótica Artocarpus heterophyllus (Moraceae) invade a Mata Atlântica brasileira

2010·4.8 MB·English

by  Rodolfo Cesar Real De Abreu

Rodriguésia61(4):677-688.2010
http://rodriguesia.jbrj.gov.br
Exotic tree Artocarpus heterophyllus (Moraceae) invades
the Brazilian Atlantic Rainforest
ÁrvoreexóticaArtocarpusheterophyllus (Moraceae)invadeaMataAtlânticabrasileira
Rodolfo CesarRealdeAbreu'2&PabloJoséFranciscoPena Rodrigues'
Abstract
ArtocarpusheterophyllusLam.(jackfhiit)wasmanmadeintroducedinTijucaNationalPark(TNP)inthemid-
1800s. Native from Southcast Asia, nowadays denscly colonizes the TNP. Herc we analyzc somejackfruit
populationparametersthatprobablyallowedtlielocalcolonization followedbysuccessful invasionofthencw
habitat.Basedon20sampleplots,randomlyplacedinsidc5sitescolonizcdbyjackfruit,wedescribcdsubpopulations
hierarchical size diameter structures. Gini’s coefficient values tumcd around 0.64 and Lorenz’s asymmetry
coefficientaround 1.03,indicatingthatsizehierarchieswereverysimilarbetweensites.Theftvesiteswerealso
compared and do not diffcr based onjackfruit basal area, numberofspecies, Shannon’s index and canopy
openness.Jackfruitmay,thcrcfore,beconsideredinvasiveinallthesesitesandthelowtreediversityvalues(H’
overallmean0.74)alsoindicatesthatthisexoticspecicslocallyexcludesnativeones.
Kcywords:jackfruit,invasivespecies,biologicalinvasion,populationstrueture,TijucaNationalPark,Brazil.
Resumo
ArtocarpusheterophyllusLam.(Jaqueira)foiintroduzidapelohomemnoParqueNacionaldaTijuca(PNT),em
meadosdoséculoXVIII.NativadosudestedaÁsia,estaespécieexóticahojecolonizadensamenteoPNT.Neste
estudo analisamos alguns parâmetros populacionais dajaqueira que provavelmente permitiram a colonização
local cconseqücntcmentea invasão bem sucedidadestenovohabitat. A partirdaamostragemde20parcelas
alocadas aleatoriamentecm cinco sítioscolonizadosporjaqueiras, descrevemos asestruturasde diâmetrodas
subpopulaçóes. OscoeficientesdeGinioscilaramem tomo0,64eosde Lorenz 1,03, indicandohierarquiade
tamanhossemelhanteentreasamostras.Oscincosítiostambémforamcomparadosenãodiferiramquantoàárea
basaldejaqueiras,riquezatotaldeespécies,índicedediversidadedeShannoneaberturadodossel.Ajaqueira
pode,portanto,serconsideradainvasoraemtodosestessítioseosbaixosvaloresdediversidadedeárvores(H’
emtomode0,74)tambémpodemindicarqueestaespécieexóticaexclui localmenteasespéciesnativas.
Palavras-chave:jaqueiras,espéciesinvasoras,invasãobiológica,estruturadepopulações,ParqueNacional
daTijuca, Brasil.
Introduction al. 2001). Artificial introduetionscan bcextrcmcly
harmful to local ecosystems, causing negative and
Human-mediatcdchangesinnaturalecosystems
have caused profound biota modifications, and evenunrecogni/edimpacts(Williamson 1996).Mainly
becausesomeexoticspecicsareabletochangclocal
prominent changes in geographic distributions of
plantsandanimais(Cronk&Fuller1995;Williamson ccosystemfunctioning(Kourtevetal. 1999;Zavaleta
1996;Pimentcletal. 2001). Speciesnaturallycross etal. 2001) and biodiversity patterns (Stohlgren et
geographical barriers, but human activities such as al.2001).
Invasiveexoticspecics(IES)alsomaycause
intemational commcrce and intercontinental travei
have strongly contributed to accelerate these nativespeciesextinctions,insomecasescomparablc
transpositions (Richardson etal. 2000; Pimentcl et togeologicalmass-cxtinctions(Casscyetal. 2005),
’InstitutodcPesquisasJardimBotânicodoRiodcJaneiro,R.PachecoLcâo915.22460-030.RiodeJaneiro.RJ,Brasil.Autorparacontspondíncia:[email protected]
turrenladress:PPG-SEA-CentrodeRecurso.HídricoseEcologiaAplicadaCRHEA/EESC/USP.Av.TrabalhadorSâoCarlcnsc400.13566-590,SSoCario»,SP.
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leading to biota homogenization (McKinney & introduced(Atalaetal. 1966;Bandeira1993;Dean
Lockwood 1999). World’s biodiversity reduction 2002).ThisprogramwasthefirstBrazilianinitiative
is straightly related to habitat loss and invasion to recover a forest ecosystem previously
(Simberloff2003),bothimpactsfullyexperienced devastated by sugarcane and coffee cycles (Conti
by Brazilian ecosystems (e.g., Pivelloetal. 1999; et al. 2008). But the widespread information that
Scariot 2001). However, the magnitude of the the whole TNP (4.000 ha) was recovered by that
problems generated by biological invasions will restoration initiative it is not true. Environmental
vary depending on the exotic species introduced, historydataofRiodeJaneiroAtlanticforestshowed
onthelengthoftimesinceitsintroductionandthe thatthesereforestationwasresponsibleby 100,000
susceptibility of the environment to invasion seedlingsplanted,covering 170ha(Oliveira2007).
(Williamson 1996). Infactremainingforestfragments,onhigherareas
In Brazil, only a few studies describe IES oftheTijucaforest,probablycouldactaspropagule
biology in new habitats, especially trees. Mostly sources colonizing the deforested areas and also
invasionprocessdetectiononlyoccursafterhigh enriching the restored sites. Nowadays, in some
dominance of the invader or another advanced areas at TNP A. heterophyllus is dominant in
impact on local community can be recognized density and biomass (Abreu & Rodrigues 2005,
(Williamson 1996). Since an exotic species is Cunha et al. 2006). These populations may have
detected colonizing natural habitats, population originated from some individuais that were
structure may reveal the capacity of population historically introduced in the above-mentioned
expanding at present, once this structure can reforestation, from which the exotic species has
indicatedemographic futureofspecies (Bruna& reached the status ofinvasive, since it appears to
Kress 2002). In this aspect, ancillary tools that haveexpandednaturally.
describe the degree of inequality in structured Furthermore, we compared the population
populations have been widely used in the size structures of the invasive tree A.
dpoepsuclraitpitoinosn(Woefinheirer&arScohlibersigof198s4i;zeSsolwbiritghi&n hweetetreosptheydlltuhseinfoflilveowsiintegs tofwoTNpPr.ediSpcetciiofnisc:all(y1,)
Solbrig 1984).Amongthese,wehighlighttheGini's population size structures change among sites,
coefficient and the Lorenz’s curve. Gini’s suggestingthatsomesitesaremoreinvasiblethan
coefficient has been used in biological studies others; (2) diversity indexes decrease in places
generally to describe size hierarchies and/or occupied by jackfruits. In testing these
i1n9t8r5a,spe1c9i8f6i;ccDoamm&pgetaiatridon&ofWaeninnuearlp2l0a0n0t)s,(sWyesitneemr hofypTotNhPesseusf,feoruerdgmoaaljoirstiomdpeatcetrsmdiuneewtohibciholpolgaicceasl
crops (Sadras Bongiovanni 2004) and tree invasion, and to setpriority sites to management
populations on natural habitats (Rouvinen & and control ofthis invasive species.
Kuuluvainen 2005; Bagchi 2007). On the other
hand, theLorenzcurvecanbeabetterdescriptor Material and Methods
than skewness, generally used to describe
population structures (Weiner & Solbrig 1984; Study area
Damgaard & Weiner 2000). Furthermore, a Tijuca National Park (TNP) is one of the
population’s stage structure can be the basis for largest urban forests in the world, and is widely
immediatemanagementdecisions(Bruna& Kress usedasarecreationalsite(22°55’-23°00’S&43°11’-
2002), specially related to invasive species with 43°19’W). Three fragments constitute TNP.
long lifecycles and increasing populations. subdividedbyfoursectorswith4.000haaltogether
ThejackfruitArtocarpusheterophyllusLam. inside Tijuca Massif, compound by innumerous
(Moraceae)isanexoticspeciesnativefromSoutheast mountains that reaches until 1021 a.s.l. (IBAMA
Asia, which now occurs widelyin reserves ofthe 2004) situated at the Brazilian second-largest
Atlantic Forest in Rio de Janeiro, Brazilian economicandpopulationalcenter(IBGE2006).This
Southeast. Probably the introduction ofjackfruit factmaycauseimpactssuchasuncontrolledurban
trees into some areas of the lowest part and the growth in the boundary regions (Coelho-Netto et
middle slopes ofthe Tijuca National Park (TNP) al. 2007), atmospheric pollution (Azevedo et al.
beganin 1862withareforestationprogram,inwhich 1999)andthepresenceofnon-nativefauna(Cunha
bothnativeandexoticspeciesweresimultaneously etal. 2006)andflora(Bandeira 1993).
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Jackfwit Invades the Atlantic Rainforest
Theannual precipitation varies from 1300 asynchronously during the entire year, with a
mm to slightly more than 3000 mm, with large proportion prcferentially fruiting in
abundant precipitation also in winter (Mattos summer (December-February). The fruits are
etal. 1970)andtorrentialrainsinthehotsummer consumed by humans and by other mammals
months (December-February) (Coelho-Nettoet suchasrodentsandprimates(Cunhaetal. 2006).
al. 2007). Lithosols, cambisols and latosols Fieldobservationsindicated ahighgermination
predominateintheupperpart, andpodzolicsoils rate ofseeds in edge areas orwhere the species
in thelowerpart (Coelho-Nettoetal. 2007). alrcady occurred (R.R. Oliveira, personal
TheTNPvegetationistypicaloftropicalrain communication). Prcsently, its distribution
forests, characterized by big trees, palms, fems, includes nearly theentireterritoryofBrazil in a
epiphytes,andlianas.Inthematureforest,emergent fairly wide range of habitats, colonizing open
trees reach 25 m height and canopy strata mean and forested areas associated with human-
heightisabout20m(Atalaetal. 1966). Withinthe impactedcnvironments(Carauta& Diaz2002).
TNP, there is amosaic ofvegetation ranging from In RiodeJaneiro,itdenselycolonizesedgeareas
undisturbed forests to patches totally invaded by ofthe Atlantic Forest, and is found in Reserves
African grasses. Notwithstanding, the most such as the Tijuca National Park and the Poço
representativefamiliesinthatstudywereMyrtaceae, das Antas, União and Tinguá biological
Lauraceac,Leguminosae,Rubiaceae,Euphorbiaceae, reserves (Abreu & Rodrigues 2005).
McliaceaeandAnnonaceae.Themorerepresentative On the native habitat (Southeast Asia), A.
spccieswereCoussareanodosa(Benth.)Müll.Arg„ heterophyllus is typical of advanced succcssional
Geonoma schottiana Mart., Myrcia rostrata DC, stages,andgrowsnaturallyuptoanaltitudeof1300
Myrcia laxiflora Cambess., Allophylus edulis (A. m in the sub-canopy ofthe MonodominantHumid
St.-Hil.,Cambess.&A.Juss.)Radlk., Cryptocarya Tropical Forest.The species isconsidered rare(up
inoscliata Nees & C. Mart., Tetrorchidium aff. to 1 individual perha) in this environment, where
rubrivenium Poepp., Eugenia prasina O. Berg, few species dominate the different forest strata
PausandramegalophyllaMüll.Arg.andTrigynaea (Chittibabu & Parthasarathy 2000; Bhuyan et al.
sp.Schltdl.(Oliveiraetal. 1995). 2003). It is shade-tolerant, and germinates best in
clearings;thefruitsmaturebetwcenJulyandAugust,
The studied species andthe seedsvary in sizeandweight(1.5 to 14g).
ArtocarpusheterophyllusLam. isaspecies Theirdispersersincluderodents,monkeysandwild
originated from Southeast Asia (Chaves et al. pigs(Khan2004). Theseedshavearelativelyhigh
1967). In the year 1682, in the State of Bahia, rateofpredation(Sodhietal.2003)andgerminatein
otrhiegrien.weTrheealfirrcsatdrye1co1rjdacokffrauislhtirpemesenotfutonkBnraozwinl wthheernain8y5se%asoonf,dtuhreinagntnhueaslumpmreecri(pMiatyat-iSoenptoccmcbucrrs)
dates from January 1683 (Ferrão 1993). The (Chittibabu&Parthasarathy2000;Bhuyanetal.2003).
recently arrived seeds and seedlings were sent Inthismanner,theseedsgerminateandtheseedlings
to a nursery in Bahia State to bc acclimated, establishthemselves,becomingablctoeasilytolcrat&c
and were later distributed to other regions of the December-February dry season (Chittibabu
thecountry(Dean2002).In 1803inRiodeJaneiro Parthasarathy2000;Bhuyanetal. 2003).
municipality, naturalized adult individuais
apparentlyalrcadyexisted (Chavesetal. 1967). Sites descriptions
Thejackfruitisalargetree,reachingaheight Five study sites weresamplcdatTNP.Thcy
of more than 10 meters; the trunk may exceed 1 have similar disturbancc histories, because
meterindiameter.Itismonoeciousandcauliflorous comprisedareaswhichwereatonctimedeforested
(Chavesetal. 1967;Craneetal. 2002).Thespccies forsugarcancand/orcoffeecultivation,andwhere
can be pollinated by insects or wind, tolerates thejackfruittreeswerelaterintrodueedandnaturally
climate stresses well, and in ideal orchard disseminated. Random sampling mcthods were
conditions it can fruit in three or four years used to demarcat&e plot locations at each site
(Crane etal. 2002). In Brazil, awell-developed (Muellcr-Dumbois Ellcnberg 1974).
jackfruit tree can produce up to 100 fruits per Site 1-Locatcdinthe“A”sectorofTNP,afragment
year, some wcighing more than 30 kg (Chaves with approximatcly 1,300 ha. This site is locally
et al. 1967). Individuais of this species fruit known as “Represa dos Ciganos”. Six plots were
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placedatNorthernexposed slopes. This fragment for unequal Ns; p < 0.05) were used to assess
hasawellstructuredforestwithtalltreesandthere whether the parameters analyzed (number of
werethetallestjackfruittrees. individuais,totalbasalarea,basalareaofjackfruits)
Site 2-Locallyknownas “HortoFlorestal”,that differed signiflcantly among sites, because any
site belongs tothe “B” sectorofTNP, the bigger difference found may be indicative of different
fragmentwithapproximately 1,660ha.Twoplots processes and patterns experienced by each
were placed close to the well preserved forest subpopulation.Thesite2 wasnotconsidereddue
remnantknownas“MatadoPai Ricardo”,located tosmallnumberofreplicates(2).
atSouthernslopenearaboundaryregion. Atthat To test the hypothesis that (1) population
place,jackfruit densely colonized roadsides and sizestructurechangeamong sites, suggestingthat
it is believed that jackfruit root systems can some sites are more severely invadedthan others,
prevent sliding ofslopes. we performed trunk diameters structure from the
Site3-Thissiteislocatedinthe“C”sector,which five sites and made comparisons between than
isaforestfragmentwithhighhills,coveringabout throughfrequencydistributions,measuresforsize
230 ha. Site 3 comprises fourplotsplacedonthe inequalities in population structure, and spatial
Southernslopes.Twofromtheseplotswereplaced distribution at plot scales.
intheriparianarea. Attemptstoconfirmtheageoftheseindividuais
Site4-Itisalsolocatedinthe“C”sector,butplots bytheirwoodanatomywereunsuccessful.sincethis
were established on the Northern slopes. speciesdoesnotformgrowthrings.
Site5-Thissiteislocallyknownas“ParqueLage”,
whichwasasugarcaneandcoffeefarminthepast, Frequency distribution
atpresentcoveredbysecondaryforestandplanted Diameterclasses were established according
totrhfeeeTsoN.tPSh.eirnTcaheree2a0vse0go4eftitTawtNaiPosnebsumttbrueecxdtodutreiedcitssopvetechrieye“ssBio"mciscleuacrrtotaort dt(oeu.rgmi.on,rgppthrhoeelsopeginilcoctealsotfcuhdaryreatpcortodederufsicnotebisvoeenrtvsoetgdreunicenttuitrcheesstfaaigeneldsd
the bottom regions. Fourplots were located close characteristics of the crown and/or trunk). A
to a trackside used by park visitors (track to frequencyhistogramweredeveloped,wheretheclass
“Corcovado”), exposedtoSoutherndirection. JUVENILES 1 was composed ofsmall sized trees
DBH
with rangingfrom 1 to4.9cm;JUVENILES2
Structures of trunk diameters wascomposedofsmall-tomedium-sizedtreeswith
(10 mInrtadhieuyse;ar3124004m,22e0accihr)cuwlearrepersemtanteonstuprlvoetys dDeBveHlorpaendg.inTghefrionmdi5vitdoua9i.s9ocfmclaansdsJthUeVcErNoIwnLEliSttl3e
jackfruit populations at TNP. The 20 plots were included small tomédiumtreeswith DBHranging
wdihsetrreibjuatcekdfriuniftipveopsuiltaestisocnasttweerredetrhercoorudgehdo.utTNP, dfervoemlo10petdo.14T.h9ecmcl,asstsillofwitPhRtEhe-cRrEoPwRnOreDlaUtCivTelIyVliEttSle
Allindividuaisoftheplantcommunitywitha included individuais from 15 to 24.9 cm DBH,
mdiaarmkeetderanatdbmreeaassturheedig.htIn(tDhBeHy)ea>r520c0m9,wealrletthhoesne cwiotmhpoaserdeloaftliavregleyimfumlaltucrreo(wnno.n-Trehperodruecptriovdeu)cttrieevse.
wcianildtcihuvlDiadBtueaHiisn>dwie1vricedmuaawlgeabrianesaimlnecaalrsueudaresed(diBnaAnt)hdewisenudruivsveieydd.uatTihose Di1nBadinHvdifd2ur.aoTimsh2ew5ecrtloeas4ds4i.Av9iDdcUemdL,iTwnitStoht1haiesnccmlloauosdsteehdstAtrruDeneUks.LwTTihtSeh
f(coirrmcuDlleBaa:rHeaS)=;7tn.r=2;3.w1h4eraen:drS==rianddiiuvsi(druaadliubsas=ahlalafreoaf celqausaslAoDrUgLrTeaSte2ritnhcalnud4e5dicnmdi,vigdeunaeirsawliltyhwaidtihambeatrekr
the measured). Individuais with visibly fissureson the trunk.
interlinked multiple trunks were considered as a
single individual; in these cases, each trunk was Population inequality measures
measured separately and their BA were summed,
cadainladcmuefltrTaeohtrmeed>tbhf1aercsorammelswutaalhrtseienoaDgbsBtbHaoaisfsn.aealdl.Olanrieena-dWaivasiyidnugAalNiesOviVwretAurasel cWLuoerrivenenseS,zriGzsi2en0aii0’sn0sey)qcmuomaaelefniftdtirycyitwehnaectisore(afWrsfeesiisecnpsieesecrnettd&iuv(SseDoialncbmgorgnLiafgoiar1dre9end8nz4c')&es,
followed by an a posteriori Tukey’s test (HSD rinatnekrevadlsa.ccIonrtdhiengLotroetnhze’irs csiuzreve(,DiBndHiviindutahiisscwaesree)
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Jackfrult Invades theAtlantic Ralnforest
/ > 1 aggregateddistribution,andIm < 1 uniform
distribution, and a measurement ofthe degree of
aggregation, whereas other measurements of
dispersion test only one of these patterns. This
index has desirable statistical properties: it has a
knownsamplingdistribution,andisnotinfluenced
by sample size. To test the significance of the
values ofIm of 1.0, the chi-square test was used:
X2(df=Q-l)=(Q-l)s2/p(Souza&Martins2002
cipudDavid & Moore 1954), where Q is the total
number of plots, and s2 and p are, respectively,
the variance and mean ofthe numberofjackfruit
trees perplot.
DBH(cm) Richness, biodiversity
Figure1—Diameterhistogramofjackfruitsubpopulations and canopy structure
atTijucaNational Park, RiodeJaneiro, in2009(DBH 2 Communitysampledtreeswereclassifiedin
lcm).Theverticallinesindicateonestandarddeviationfor morphospecies. From these data, PC-ORD 5.19
each class. (McCune & Mefford 1999) was used tocalculate
richness and Shannon-Weiner biodiversity index
andcumulativefractionofthepopulationsizewas ineveryplot.
plotted against cumulative fraction of total To describe canopy structure, we calculate
populationsize. Ifindividuaiscontribute tooverall canopyopenness aftertakingdigital pictures with
populationsizeinproportiontotheirownsize,then aNikonCoolpix990,afisheyelens(NikonFC-08)
astraightlineofequalityisexpected.Hierarchydegree and a tripod one meter high. Pictures were taken
canbesummarizedbyGini’scoefficientrepresented accordingtoterrainslopeinthecenterofplotsduring
bythedifferencebetweenequalitylineandLorenz s cloudyhomogeneousskiesfrom2007summer.To
curve(Solbrig&Solbrig 1984;Damgaard&Weiner analyzepicturesweusedWINSCANOPY2003.din
2000). Gini’s coefficients calculated should be anautomaticthresholdmanner(Regent2003).
multiplied with n/fn-l) to the estimates become Comparisons among sites were done after
unbiased (Weiner 1985). Lorenz’s asymmetry normalitytests.Sincevariablesdonotfittonormal
coefficient (S) expresses internai asymmetry from distribution,Kruskal-Wallistestswereapplied.Site
Lorenz’s curve to the axis of symmetry. S values 2 was not included in the analyses due to small
from 1 indicatesasymmetricalLorenz scurve,S<1 numberofreplicates(2).
indicates asymmetry to the left and S>1 right
asymmetry(Damgaard&Weiner2000).Confidence Comparisons with native species
intervals(p=0.05)fortheestimatesofGandSfora Inordertodetectpopulationpatternsofnative
sample were obtained with 1,000 repetitions in a speciestocomparetothepopulationsofjackfruits,
bootstrappingprocedure(seeDixonet.cil 1987).To secondary data from phytossociological samples
G calculations and comparisons between sites we inareasoftheAtlanticForestwere used, in which
usedWINGINI 1.0(Santos 1996).ToScalculations onlythosespecieswiththehighestvaluesofbasal
andconfidenceintervalsweusedatrialversionfrom areawereselccted.
MATHEMATICA 7.0 with a notebook developed
byDaamgard&Weiner(2000). Results
Withintheplots,thesubpopulationscontained
odibenstdeeerrxmvSTieaphlndaleetbdí2yau0slsSpiaosntduagiizmasMapltol&rreiidsbMipiuasltrtoattit’riossinnbiswunedt(eri2xeo0.n02Tc)oop,nacstatihltdeceeuMrrloenardti.eswtiaAhtissas Jh(toiUDg1VBh0EHr0Ne%l>1atoL5ifEvcteShmde)e1ni(wsnideDtirBiveeiHsdsuo1aaftimjospa4cli.kne9fdarculimiontc)sa2,la0inr0tad9yn,.5gA6ia4nntgodbtifatmglrhgooecfmrm41te2r0ae6e%n2s
Iinnddiecxat(eIdm)bygitvheesvatlhueest:yIpme=of1 rsapantdiaolmddiissttrriibbuuttiioonn,, bwaistahlaaprperaoxoifmjaatceklfryu2i8ts±w1a1str1e.e7s1p2l±ot01.(7T7ab3. 1)2.plot1,
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The structure of the jackfruit population in changestotheecosystem.includingindirecthabitat
TNPin2009(diametershistogram)showedtheright- destruction,facilitatingtheirowndominanceover
asymmetrical log-linear distribution pattern, thenativespecies(Fine2002).Jackfruitnowadays
knownasthe“reverscJ”(y=182.4e’05138x;R:=0.72). denselycolonizesmanysitesatTijucaNationalPark
The largest proportion of the individuais (70%) (TNP), anurban and permanentlydisturbed forest
rweeprreodiuncttihveefairdsutltssizceorclraesssp,onadneddthtoe 9cl%assoefstohef sreumcnhaanst.frIangmaeddnittaitoino,niamnpdachtasbiotnatnlaotsusr,almahaybiftaavios,r
individuais ofthe population.
Thedensestsubpopulationscontainedmany the dissemination ofthis exotic species (Davis et
immature and few adults. However, as expected. al. 2000; Fine 2002). In this way, the successful
adults were responsible forthe greaterproportion TmaNnP-mediated introduction ofjackfruits into the
ofthebasalarea,whichweredistributedunequally (Atala etal. 1966; Bandeira 1993) after 150
betweenadultsandimmatures(Tab. 2). years resulted in an effectivehabitatinvasionthat
MaximumDBHrangedfrom80.9insite 1 to probably represents the most massive invasion
108.7insite3.GinPscoefficientswereallnear0.65 promotedby atree speciesat Atlantic Rainforest.
andLorenz’sasymmetrycoefficientswereallnear1 OtherBrazilian Reservesthathaveedgesites
(Tab. 3). These values indicate high similarity in colonized by jackfruit showed an increased basal
size hierarchies and no sample asymmetry were areawrhencomparedtootheredgesites(Rodrigues
observed between smaller and bigger individuais 2004).Thisfactmayindicatethatevenpopulations
in a single site. No differences were observed of native species with larger basal areas in local
between sites. Analyzing theconfidence intervals communitiesareincapabletocolonizethelocalityin
of the coefficients, all sites had the same size the same manner as the jackfruits. The jackfruit
hierarchies. In the landscape scale, Artocarpus colonization success as well as the dominance in
heterophyllus showed clumped pattems, resulting density and biomass (Abreu & Rodrigues 2005;
inaMorisita’sDispersionIndexvalueof1.11 Cunhaetal.2006)canbealsofavoredbypopulation
The sites were also very similar in terms of pattemssuchasclumpeddistribution,dispersaiway
basal area (BA), richness (S), Shannon-Wiener (barochory),andbytheprobablyescapefromnatural
diversityindex(H’)andcanopyopenness(Tab.4). enemies(Sax&Brown2000;Keane&Crawley2002).
Meanbasalareaswereabout 1.71 (±0.77)nrplot The time elapsed since the introduction of the
mean richness was 6 (+ 4) species plot1, H’ was speciesbyplantinginalocalitymayalsocontribute
about0.74andmeancanopyopennesswas6.16%. for species dissemination (Williamson 1996).
Comparing the basal area (BA) of the sub- Furthermore,thejackfruitseedpredationrateishigh
populationsofA. heterophylluswiththeBAofother in preserved areas and low in forest edges, and in
speciesoftheAtlanticForest(Tab.5),theBAoccupied thelatterareasthespeciesshowsahighgermination
byjackfruitswas 10.9to98.1 timeslargerthantheBA (RR- Oliveira,personalcommunication). Inapilot
ofnativetreespeciesoftheAtlanticForest. study (unpublisheddata) we observed single plots
(2m2)havingmorethan200jackfruitseedlings.This
Discussion shows that thejackfruit can regenerate betterthan
Although exotic species may be rare in cotahnebrespeexcpileasinferdobmetchaeusAetliannttircopiRcaailnffoorreesstts..Apnladntist
uspnedciisetsurmbaeydtbreopciocmalefionrevsatssi,vienadnidsteuvrebnedcfaourseestdsrassutcihc pwliatnhtlsawrigetrhssemeadlsltseeneddtso(gDeramwisnoanteetmaolr.e20r0ea9d)i.lythan
c2c0oo0mm9mmuu(nnDiiBttyyH(>tmrelehesa.mh)a);XBBANA^,^=n=uebmnabtsoeaelracorfoejmaamocuckncfiuntpmiyien(ddnbifvyihdoautahie)srbiainsnatdlihveaicrdeouaam;imsBu,nnAiottliljqy=a(ctkbríaehseiasilhtasa'r(1)em.aM:cheoaam1np);o±NsSeumDdI==exmnceuluamsnbievaernlodyfsoitfnajdnaidcvakinfdiruudaieitvsiiianntittohhnee
ofparametersbyplot(314m').
BAtnJm2ha') BAi«M(m2ha'> BAo,heJm2ha ') N.„Jtreesha‘) N. (treesha')
TMoetaaln±SD 2.16489.+404.6787 1.65449.+3501.762 0.91552.±921.6115 34.71150+79.5 27.98+8191.5
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Table2-Jackfruitdemographicpattemsofthestudysites(mcan±standarddcviation).Sites=localities(numberofsitesinthclocality,
sne=emneutmhboedrsotfexitn)d;iIvimdmuaatius;rc%sn==inpdeirvcideunatiasgweioftnhuDmBbHerraonfgiinndgifvirdoumai5s;toB2A4.=9cbmas;aAldaurleta;s%=iBndAiv=idpuearicsewnitatgheDoBfbHagsraelaatrecrat;hAanN2O5cVmA;
=valuesofMSforOne-WayANOVAs,df= 15,p<0.05,Tukey’stestforunequalNs.Symbol(*)indicatestliatnosignificant
differenceswereobserved.
Immaturcs Adults
Sites n % n BA %BA n % n BA %BA
1(6) 23±10 71 0.28±0.14 17 10±6 29 1.38±0.82 83
2(2) 25±13 79 0.34±0.19 19 7± 1 21 1.47±0.25 81
3(4) 16±7 73 0.16±0.07 10 6±1 27 1.47±1.07 90
4(4) 15±5 61 0.19±0.11 13 9±5 39 1.27±0.72 87
5(4) 22±1 73 0.22±0.10 12 8±3 27 1.53±0.62 88
ANOVA* 82.889 0.01498 17.167 0.63681
Table3-Parametcrsanalyzedtodescribethestructurehierarchyofsizcs(DBH)inthesitesstudiedintheTijucaNationalPark,
RJ(2009).Site=studiedsite;n=numberofindividuais;mcan±SD=meanandstandarddcviationofDBH;maxDBH=greaterDB1
observedinthesite;G’=corrcctedGini’scoefTicicnt;ClG’95%=confidenceintervalforG’;S=Lorcnz’AsymmctricCoefficient;
ClS95%=confidenceintervalforS.
Site n DBH maxDBH G’ CIG’95% S Cl S 95%
1 723 6.77±11.42 80.9 0.6452 [0.6232-0.6615] 1.0346 [1.0006-1.0794]
2 1% 7.80±13.07 81.9 0.6481 [0.5931-0.6815] 1.0375 [0.9497- 1.0977]
3 205 10.41±17.39 108.7 0.6467 [0.5891-0.6780] 1.094 [0.9890-1.1470]
4 298 9.15±10.03 83.6 0.6551 [0.6328-0.6736] 0.9682 [0.9050- 1.0304]
5 404 7.46±13.51 107.7 0.6459 [0.6061-0.6753] 1.0317 [0.9843-1.0954]
Table4-Means(±SD)frompanunctcrsusedinthecomparisonsbetwccnthesites.DBH1=Basalareacomposedofjackfruits
witliDBH>1cm;DBH 10=BasalareacomposedofjackfruitswithDBH> 10cm;S=speciesrichness;H=Shannon-Weiner
biodiversityindex;Openncss=canopyopenness(%).NosignificantdilTcrcnccswereobservedbetweensites.
DBH1 DBH10 S H Openness
Site
1 1.67±0.99 1.58±0.97 4.33±2.07 0.53±0.45 5.64±0.48
2 1.78±0.00 1.70±0.02 6.00±1.41 0.72±0.44 5.88±0.35
3 1.65±0.90 1.59±0.86 4.25±3.59 0.75±0.80 6.16±0.34
4 1.64±0.88 1.59±0.87 9.75±6.24 1.18±0.92 7.66±1.33
5 1.89±0.72 1.80±0.70 5.25±1.71 0.62±0.05 5.60±0.70
Ecosystemsinvasibilitymustnotbegeneralized, attributcsoftheinvasivespeciesandthesusccptibility
bccause they are closely dependent on scale, ofthcinvadedecosystem(Williamson 1996;Heger&
vegetation type, biome, and resources availability Trepl2003).Additionally,disturbanceandpropagulc
(Stohlgrenetal. 1999).Furthermore,insomecasesit pressure are factors that deserve attention because
isnotnecessarytotheinvadertobcaggrcssivewhen although they are confused, both factors play
this species arrived to occupy a potcntial availablc importantrolesininvasionoftropicalforests(Edward
niche(Heger&Trepl2003).Jackfruitthatcolonizes &Munishi2009).
high biodiversity Atlantic Forest arcas likeTNP, at Jackfruit in his original habitat is rare,
first excludes local plant species. In this case, the experienccs a monsoon seasonally climatc
invasion success was determined by both the condition (Ayyappan & Parthasarathy 1999;
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Table5-SecondarydataextractedfromphytossociologicalsurveysinAtlanticForestareasinRiodeJaneiro.DBH=minimum
dspieacmieetserexatdroapptoeldaatsedintcoluhseicotnacrreiste;risao;uPrc(eha=)r=efseirzeencoefforroimgipnhayltossasmopcliionloggairceaalisnurhveecyt.ares;BA=basalarea(m:)occupiedbythe
Species DBH(cm) P(ha) BA(m2ha‘) Sourceandplace
ArtocarpusneterophyllusLam. 5 0.63 53.68 This study*
Artocarpus heterophyllus 10 0.63 5225 This study*
Metrodorea brevifolia Engl. 10 1 4.13 Silva&Nascimento(200
Paratecomaperoba(Record&Mell)Kuhlm. 10 1 126 TabuleirosdoNorte-R1J
Pseudopiptadenia contorta (DC.) 10 1 0.57
GRLewis&M.P.Lima
Trichiliapseudostipularis(A. Juss.) C. DC. 10 1 054
HyeronimaalclwmeoidesAllemão 10 0.6 4.26 Morenoetal. 2003*
Pseudopiptadenia contorta 10 0.6 2.15 RegiãodoImbé-RJ
&
Rustiaformosa(Cham. Schltdl.exDC.)Klotzsch 10 0.6 213
Virolaoleifera(Schott)A.C.Sm. 10 0.6 210
Actinostemonverticillatus(Klotzsch)Baill. 10 0.6 1.60 Morenoetal. 2003*
MabeafistuliferaMart. 10 0.6 157 RegiãodoImbé-RJ
Maytenuscommuta Reissek 10 0.6 153
Vochysia oppugnata (Velloso)Warm. 10 0.6 1.47
Cupania emarginata Cambess. 10 0.4 4.81 Rodrigues2004*
Ecclinusa ramiflora Mart. 10 0.4 3.08 REBIOUnião-RJ
Ficusgomelleira Kunth&C.D. Bouché 10 0.4 2.93
Pradosia kuhlmannii Toledo 10 0.4 2.87
Virola bicuhyba (Schottex Spreng.) Warb. 10 0.4 2.78
Alchornea triplinervia (Spreng.) Miill.Arg. 5
1 5.1 Guedes-Brunietal. 1997
Cabraleacanjerana(Vell.)Mart. 5 1 12 MacaédeCima-RJ
Euterpe edulis Mart. 5
1 1.9
Mollinedia gilgiana Perkins 5 1 15
Myrcia pubipetala Miq. 5
1 1.4
Casearia obliqua Spreng. 5 1 1.6 Pessoaetal. 1997
Crotonfloribundus Spreng. 5 1 2.6 MacaédeCima-RJ
Euterpe edulis 5 1 1.9
Nepheleasetosa(Kaulf.)R.M.Tryon 5 1 12
Tibouchina scrobiculata Cogn. 5 1 3.9
* extrapolatedvaluesforhectares forcomparisonpurposesonly. specieswerenotobservedin acontinuoushectare.
pPrarotcheasssaersat(hKyh1a9n992)0a0n4d).exOhitbhietrswyinsceh,roinnictihteyiBnrafrzuiiltiianng iwnassoamlseoionbvsaedrevdedenivniTrNonPmtehnattsja(cJkafkrouibtssedtoaln.o2t0e0x1h)i.biItt
Atlantic Forest the absence ofa marked dry season synchronicity in relation to the fruiting period, even
ce(oxMnpatetictntouesededtthraaelt.perx1oo9dtu7ic0ct)sigwonirtomhwawfyaatsbteeerrfsaaunvpdoprrleeydp.yreoIandruf-carecotb,uentidtteiars cdtioifnmfseeisndeoenfrtitinhngediytvehiaedrur.aeTipshr,iosedvuisecinancngeopimgehmabkoornats,psafutrmutimetemarti.ndiTsfhpfaeectrieiensst.
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of the family Moraceae (J.P.P. Carauta, personal hierarchies, richness, biodiversity and canopy
comunication),asoftenobservedinfigspecies(Romo openness,itisprobablethatjackfruitispotentiallyan
1996;Korineetal.2000;Wcndelnetui2000). invasive species at TNP, even in places with small
Asymmetriccompetition incrcases variation in populations at present. All data showed that the
growthratesbetweendominantandsuppressedplants, BrazilianAtlanticRainforestisanewhabitatthatallows
its onset promotes size inequality, and this effect is the jackfruit to proliferate and dominate the plant
heightened by time (Crawley 1997). Even with an community. So any place wherejackfruit colonizes
uncertainelapsedtimeafterspeciesfirstestablishment nativeforestsinthisecologicalrcgioncanbeapriority
at each site, 150 years after introduction in TNP, areaformanagement,eradicationbeingrccommended
populationcharacteristicsgivetothejackfruitthesame atthefirstdetection.
sizehierarchyandavetysimilarpopulationstructureat
Acknowledgments
different sites, showing that the biological invasion
processis verysimilarinallsites.Jackfruitshoweda We are very grateful to ICMBio (Instituto
populationstructuretypicalfromshade-tolerantspecies, Chico Mendes) from Tijuca National Park and
withmanyindividuaisinthefirstsizeclass(JUVENILE UNIRIO (Universidade Federal doEstadodo Rio
1).Together,thebiologyofaspeciesandstructuresof deJaneiro)forlogisticalfacilities;PETROBRASby
diameterscanprovidevaluableinformationaboutthe Financial support (Research funding to Programa
successional process undergone by a subpopulation Mata Atlântica/PabloJ.F.P. Rodrigues); Students
(White 1980). Different sizeclasses may compose a thathelpusatfieldwork, twoanonymousreferees
stablepopulation,andtheproportionsoftheseclasses andtoMSc.TalitaSoaresReis,Dra.GiseldaDurigan
willvaryinaccordancetotheabioticandbioticfactors andDr.AndreMantovani Oliveira forsuggestions
those population individuais and their ancestors on the manuscript.
experienccd historically (Sarukhán 1980; Hutchings
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