Floristics and life-forms along a topographic gradient, central-western Ceará, Brazil / Florística e formas de vida ao longo de um gradiente topográfico no centro-oeste do estado do Ceará, Brasil

2011·8.5 MB·English

by  Francisca Soares De Araújo

,
Rodriguèsia62(2):341-366.201I
http.'/Zrodriguesia.jbrj.gov.br
Floristics and life-forms along a topographic gradient,
central-western Ceará, Brazil
Florísticaeformasdevidaaolongodeumgradientetopográficonocentro-oeste
doestadodoCeará,Brasil
FranciscaSoaresdeAraújo' 2 RafaelCarvalhoda Costa1 Jacira RabeloLima'
, ,
Sandra Freitasde Vasconcelos' Luciana CoeGirão', Melissa SouzaSobrinho',
,
Morgana MariaArcanjoBruno1 SarahSuedGomesdeSouza1 EdsonPaulaNunes1
, & , ,
MariaAngélicaFigueiredo',Luiz Wilson Lima-Verde' MariaIracemaBezerraLoiola'
Abstract
To tcst whether thc Hora is organized in discrete or continuous units along a topographic gradient, thrce
physiognomicswcrcasscsscdondiffcrcntsoilclassesinasemi-aridregionofnortheastemBrazil:caatinga
(xericshrubland)ataltitudesfrom300to500m,deciduousforestataltitudesfrom500to700mandcarrasco
(dcciduousshrubland)at 700m. Ineachphysiognomyaspccies inventorywascarricdout,andplantswcrc
classifiedaccordingtolifc-andgrowlh-forms.Spccicsrichncsswashighcrinthcdcciduousforest(250)than
inthccarrasco(136)andcaatinga(137).Thccaatingasharedonlyafewspccicswitlithccarrasco(6spccies)
andthcdcciduousforest(18spccics).Thchighestspccicsovcrlapwasbetwccnthcdcciduous forestandthc
carrasco(62spccics).Onehundredandfourspcciesoccurredonlyinthccaatinga, 161 onlyinthcdeciduous
forestand59onlyinthccarrasco.Woodyspccicspredominatedinphysiognomicsonsedimentarysoilswith
latosolandarcnosol: 124spccicsoccurredinthcdcciduousforestand68inthccarrasco. Inthccaatingaon
crystallinebasement rclicfwith predominanceofplanosol, herbs showcd thc highestspccics richncss(69).
ComparingthcbiologicalspcctrumofBrazilianplantlife-forms,thccaatingastoodoutwithhighcrproportion
ofthcrophytcsand chamacphytcs. Considering thc Hora ofthc threephytophysiognomics studicd herc, wc
canaffirmlhatthccaatingaisadiscrctcfloristicunit.
Kcywords: vegetationclassification,biological spcctrum,growth-form,phytoclimatc,plantcommunity.
Resumo
Paraverificarscacomposiçãoflorísticaconstituiunidadesdiscretasoucontínuasao longodcumgradiente
topográfico foramanalisadastrês fitofisionomias(caatingasobrealtitudesdc300a 500m, florestadecídua
sobrealtitudesdc500a700mccarrascosobreatitudesdc700m)sobreclassesdcsolosdistintasnosemi-
áridosetentrional doNordestedo Brasil. Emcada fisionomia foi realizadoo levantamentodasespécies, as
quais foram classificadas em formas dc vida cdccrescimento. A riquezadc espécies foi maiorna floresta
decídua (250) do que no carrasco (136) c na caatinga (137). A caatinga apresentou poucas espécies cm
comumcomasfitofisionomiasdccarrascooudcflorestadecídua(6c 18espécies).Amaiorsobreposiçãodc
espécies ocorreu entre a floresta decídua c ocarrasco, 62 espécies. Foram exclusivas da caatinga, floresta
decídua c do carrasco, 104, 161 c 59 espécies, respectivamente. Quanto ás formas dc crescimento, nas
fisionomias sobre relevo sedimentarcom Latossolo c Arcnosolo predominaram espécies lenhosas: 124 na
florestadecíduac68nocarrasco.Nacaatingasobrerelevodoembasamentocristalinocompredominânciadc
Planossolo,amaiorriquezadcespécies(69)foidcervas.Naanálisecomparativadoespectrobiológicocom
outrasformaçõesbrasileiras,odccaatingascdestacoudosdemais,constituindoumaunidadeindividualizada
pela maiorproporçáodcterófitosccaméfitos. Em relaçãoà floradastrêsfisionomias, objetodesteestudo,
pode-seafirmarqueadacaatingarepresentaumaunidadediscreta.
Palavras-chave:classificaçãodcvegetação;espectrobiológico;forniadccrescimento;filoclima,comunidadevegetal.
'UniversidadeFederaldoCeará,Depio.Biologia.CentrodcOíncim,bloco906.CampusdoPiei.60455-760.Fortaleza.(P.Brazil.
;Corrcpondingauthor:tchcscawufc.br
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Introduction
indifferentdirectionsrefiectenvironmentaleffects,
At a global scale, the main environmental especially related to climate, on plant adaptations
variables used to classify vegetation are climate observedinacommunity(Raunkiaer1934).Hence,
zones. A group of similar vegetation types that whereas the growth-form classification is used to
occurinsimilarclimatezonesindifferentcontinents characterize community structure (because some
isknownasavegetation-typeorbiome(Whittaker formsaredominantormoreconspicuous),thelife-
1975, 1978a,b;Box&Fujiwara2005). formspectrumdescribesenvironmentaladaptations
Changes in topography or microclimate can of the species that compose that community
affect the biology of the vegetation, leading to (Whittaker 1975;Raunkiaer 1934).Indirectly,this
particularitiesthatcan be detectedonly ata local system provides information on local seasonality.
scale (Spellerberg & Sawyer 1999). Gradual AccordingtoWhittaker(1975),life-formsarenota
changes in climate related to topography or to structural attribute, but a floristic attribute: when
distancefromtheocean,atasmall scale,resultin thenumberofspeciesisconvertedintopercentage
continuous vegetation units, which makes a oflife-forms, this percentage would represent the
classificationbasedonfloristicattributesdifficult. spectrum of life-forms in this community or
However, when a climate variable is associated geographic area. The fact that agiven community
withdifferentsoiltypes,theregionalfloramaybe is characterizcd by particular life-forms indicates
discontinuously distributed, forming discrete speciesconvergencetowardcertainenvironmental
communities, whose limits, along a topographic conditions; and this represents a functional
gradient, can be determined by an analysis of attributeofthecommunity.
floristic composition and ofthe main growth- or In the present study, weassessed life-forms,
life-forms ofthe plant species (Whittaker 1975; growth-forms and floristic composition of three
Box&Fujiwara2005). neighboring physiognomies that occur under
Todescribccommunitytypes itisnecessary different climates, soils and topographies. These
to characterize plant forms, since physiognomy community attributes weredetermined foranarea
results from the dominant forms that compose a locatedinthesemi-aridregionofnortheastemBrazil,
community (Whittaker 1975). Classes ortypes of which comprises two geomorphological units:
plant forms are called growth-forms; this sedimentary basin andcrystallinebasement.
classification usually does not correspond to the Basedon thesedata, wetestedthefollowing
categories used by taxonomists to classify plants. predictions:i)theflorasofthetwogeomorphological
Height,woodyorherbaceoushabit,stemform,leaf unitsaredifferent,andconstitutetwodiscreteunits;
form and intensity of leaf deciduousness are ii)thelife-formspectrumvariesaccordingtoaltitude
characteristics used to define the following types and soil type, probably as a consequence of
of growth-forms (Whittaker 1975): trees, shrubs, differencesinwateravailability,resultingmainlyin
lianas, epiphytes, herbs and thallophytes. theoccurrenceofphanerophytesinthesedimentary
Instead of using a system of multiple basinandoftherophytesinthecrystallinebasement.
characteristics such as the growth-form system
proposedbyWhittaker(1975),thelife-formsystem Material and Methods
of Raunkiaer (1934) is based on a single Location and environmental
characteristic.therelationshipbetweentheposition
ofthe perennial tissue (meristem), which remains characterization of the study area
ginraocwtitvhe dsuurrifnagcet.heTwhientelrifoer-dfroyrmseoafsona,sapnedcitehse area oSfer5ra,6d4a6s Ahal,maasndNaitsurlaolcaRteesderbveetcwoeveenrstahne
coordinates 5°15’-5°00’S and 40°15'-41°00'W
crsleealpsersceitsfeiedendtbpsylaantthsseeitnetnoovffiirlvioefnelmiehfnei-tsft.oorrRmysa:ucpnhhkaairnaaecertroepr(hi1ys9tt3ei4sc),s kí()Fmicga.at1i).ngTahe(xsertiucdysharruebalahnads)twhirteheapnhyasrieoagonfom1i7.es1:
achnadmtaheeprhoyptheyste,s.hemicryptophytes, cryptophytes an ar(e2a9.o1f92%7).,93fi)ksme-as(o4n7a.l6d4e%c)idaunoduisiif)ocraesrtrawsictho
ctrheaeplrcweuoslTrealhndtte.eatwdFiorvrbeolydmsasRtpmhaeaputclntesrkauiommfap,leaoerlr,ln(ttohh1ree9m3pvaa4alt)stscepubrelnacasstrrerefdulcmoo,orrawndaeosdaf cga(reodeoemamcpoiolrfdeTpuxh1ho,e1ou.wls7io9tsgskhtihmcurfauid2alby(tl2uta0no.niatsd1rsl2)e:i%(agi)Rh).ottluihygseeucrnrlidyouesltcaaa1ltt9liei8ndn8ge)rbeiawlnsiieetfmhtaewnnaodnt
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Florísticsandlife-formsalonga topographicgradient
IMSEAR; Biodiversity inventories - Caatinga
(PROBIO-MMA)andEdital UniversaldoCNPq/
476285/2003-8. In these studies, branchcs of
angiosperms(fiveduplicates)inreproduetivephase
(flowerbuds, flowcrsand/orfruits)wcrccollcctcd
on trails and inside the best-conserved fragments
ofeachphysiognomy.Vouchcrswcredepositedin
thePriscoBezerraHcrbarium(EAC),ofUniversidade
FederaldoCeará.Botanicalidcntificationwascarried
out using analytical keys (Freire 1983; Barroso
O£23|CDCcucumirtiimunigmiauifimnl tetheal.ma1t9e7r8i,al19p8r4e,se1n9t8i6n)tahnedEbAyCcoHmpearrbiasroinumwiotrh,
Uiitiirlwltwtntul
whcn necessary, by Consulting specialists. The
Figure 1 - Location of Serra das Almas Natural cnlaamsseisfiwceartieonupudsaetdewdacsonAsPidGerIiIIng(2t0h0e9)s.ynSopencyimcys
Reserve, Cratcús, StateofCeará, Brazil.
of Missouri Botanical Garden (Tropicos.org
2009); names and/or abbreviations of spccies
authors were written in accordance with
low altitude (c. 400 m) and ii) the Meio Norte Brummitt& Powell(1992).
sedimentary basin, on its eastern margin, which
forms an asymmetric cuesta, known as Ibiapaba
Growth- and life-forms
Plateau(altitudesbctween500and700m).
The caatinga occurs in the crystallinc Eachspecicswasclassiftcdintogrowth-forms
basementcomplex, wherethedominantclassesof followingWhittaker(1975).
soils are: Solodic Planosol, Solodized Solonetz Theclassificationofeachspecicsinlife-forms
(natric Planosols)andLilhicsoils(LithicNcosols) wasdonebasedontheprotection leveiofgrowing
ataltitudesthatvary from300to500nt. tips and on lhe reduction ofthe aerial part during
In the Meio Norte sedimentary basin, on theunfavorableseason,followingRaunk&iacr(1934,
IbiapabaPlateau,theLatosoloccursontheeastern seealsoCain 1950;Mueller-Dombois Ellenberg
hogback and quartz sand (quartzarenic ncosols) 1974): therophytes (Th), cryptophytcs (Cr),
tdoheneciptdlhuaeoteutasoup,fooraennsdtLaotbcoacsucorklss,ioadnletahl(etBiretaaussdtieeslrbn1ch9to7wg2e)be.ancTk5h0oe0f chpoheanmnsicicrdroeyprphetydotpehassyt(pePhhs)a.(nHcW)ro,oopcdhhyyatmleaiseanpaahsnydatncdnsocn(acC-thwi)owocadrnycd
and700m.Thecarrascoispresentonthebackside lianas were classiftcd according to the levei of
of the plateau, on quartz sand, at altitudes of ca. reductionoftheiraerial partduringlhedry season
700 m. Weemphasize thatthe Ibiapaba Plateau (accordingtoRaunkiacr 1934).
is a ‘cuesta’, with highcr asymmetry in its
Southern part, our study arca, where there is no Data analysis
top, butan inverted V-shapcd topography where Floristic data were organized as a list with
lhe lecward on the backside exhibits a smooth families,specics,vernacularnames,liWfeeandgrowth-
forms,physiognomyandcollectors. calculutcd
declivity.
Climatedatawerenotavailable,bccausethere spcciesandfamilyrichnessforthewholedatasetand
are no metcorological or pluviometric stations by physiognomy.Tocompare the richest families
locatedon thecuesta, topand immediatebackside bctween physiognomics, weusedhistogramswith
sites on lhe Southern part ofthe Ibiapaba Plateau, lhe tenFlroircihsteisctofvacmrillaipesbcintwdeeesncepnhdyisnigogorndoemri.cswas
our study arca.
analyzed by calculating the frcquency ofspecics
and familiesinoverlappingclasses: occurrcnce in
sweavsereaTFxllthopeerrnoisfjslietovciretaclsoy:ifnrsSevaesmererpnraltveodedarmsyafArnloammgaes1m9Ne9ant9turtpaolla2nR0;e0s4leo,rnvgie-n cpa(lfchllay/apsTthiUiyíoncsggoian/o/odgcmeanycroirmdaiucsocacusaos,t,ifnoicgrnaeaasptta)ici,narasgrnarod/aldsrcccpcoshitryodiruscoitducoedsgcntiIoodomrueeioascuctshs,
CteNrPmqe/cPolEoLgDic;alInrsetsietaurtcohdporoMgirlaênmiso-dSoitSeeCmaiaátriindgoa-/ forest). Results(urc prese,nted in histograms.
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Araújo,F.S. etal.
Totestfordifferences in thecomposition of
life-formsamongphysiognomies,wecalculatedthe
life-form spectrum, which is the proportion of
speciesofeachlife-form.Wedeterminedwhichlife-
form characterized each physiognomy by
comparing our results with the normal spectrum
proposed by Raunkiaer (1934). This spectrum FAB Eli' CON MAL AST POA APO RUB MYR SAP
represents the world flora and was used here as
null hypothesis. At First, we tested fordifferences
between the obtained and the normal spectrum
usingax2test(Vieira2004).Whendifferenceswere
significam,wecalculatedtherelativecontribution
ofeach life-form’s deviation to the computed X2
statistic.Thelife-form withhighercontribution in FAB CON EUP MAL POA AST ACA APO PAS COM
each test was considered as characteristic of the
physiognomy where it occurs. Carrasco
To test for similarities with other Brazilian
vegetation types (in terms of life-forms), we
compiled studies with spectra determined for
Brazilian physiognomies (Tab. 1). We kept the
namesusedbyeachauthorforthevegetationtypes
ofeach study. To facilitate comparison, we used
only the five main life-form classes ofRaunkiaer
(1934). Hence, epiphytes and woody lianas were
included in the class phanerophytes, saprophytes
incryptophytes, andaerophytes in chamaephytes.
Wecomparedthe life-form spectrafound in Serra
das Almas Natural Reserve with those from other
studies with a detrended correspondence analysis FAB EL!P mal CON AST MYR SAP RUB PCIA APO
- DCA (Jongman etal. 1995; Batalha & Martins Families
2002); results were expressed in ordination ^Sure7~Species-richestfamiliesinthethreephysiognomies
diagrams with scores of each study and of each ofSerradasAlmasNaturalReserve,Crateús,StateofCeará,
life-form. Brazil. Abbreviations for families: FAB-Fabaccae, EUP-
Euphorbiaceae,CON-Convolvulaceae,MAL-Malvaceae,
Results AST- Asteraceae, POA-Poaceae, APO-Apocynaceae,
—
r7i2chfnaemWsislei(er2se5c0(o)Arnwdneedreex41h19i)g.shpFeearcmiiinelsit/hemesodi(ep5ch5io)sdapuenocduisesfspoefrcerisoetms. CBAoCRmAOm--eAlBicrnRaoaunmbcteiealhaceaie,acaceeeL,aaAeeM.M,-YLRa-PmMAiySar-ctPeaaacsees,aiefM,lAoPSr-AaMPca-elSapaiepg,ihnidCaaccOeeMaaee-,,
Richness valuesofthe carrasco (46and 136) and
caatinga (44 and 137) were similar to each other
and lowerthan in the deciduous forest.
Fabaceae (86 species), Euphorbiaceae (38 Familyoverlapwasaboutonethirdamongall
rsipcehceisets)plaanndtCfoanmviolilevsuliancSeearera(2d2asspeAclimeas)swNearteurtahle pthheysdieocgindoumouisesf(oFriegs.t3s)h.aHroewdetvheerh,itghheesctarnruamsbceoranodf
Reserve.Therichestfamiliesweredifferentamong tamilies,andhadthehighest(carrasco)andlowest
pfspahemycisilieysoFgiannboalamlciteehasree,e(Fwpihhgyi.sci2ho).ghnaTodhmteiheesehx(icFgeihgpe.ts2it)o.nnHuowmwabesevretrho,ef cSs(dppaeeercccriiiadeesusscoouoocsvcfaeuornrrldersaetpdt)hnwienuamsadlbleelcporihwod,yfuseaoixsuocslgounnsfolioymvrieenefsisatnme(ihFloiaigued.st(3hoF)ifi.ggT4.hh13e)e9r.
the representativeness ofsubfamilies varied, with
floristicalfinity witheachother,sincethey shared
higherrichness ofPapilionoidae in the deciduous morespecies(15%)andbothhadlowoverlapwith
forest (25 species) and ofCaesalpinioidae in the thecaatinga(1.3%overlapwithcarrascoand4.2%
caatinga (12 species) and carrasco (15 species). withdeciduousforest-Fig. 3).
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Rodrlguésla CTahb-lec1ha-mLaiefpeh-yftoersm,spPehct-raphuasneedrfooprhcyotmepsa.risonsinadetrendedcorrespondenceanalysis(DCA).Life-forms:Th-therophytes.Cr-cryptophytes,H-hemicryptophytes, Florístics
and
(622 Vegetation type Abbreviation Site Reference Th Cr H Ch Ph
):
life-forms
341 caatinga caa Sa.dasAlmas,Crateús,CE This study 47,9 1.4 6,3 18,1 26,4
3-66 carrasco can Sa.dasAlmas,Crateús,CE This study 17,2 3,4 3,4 17,9 57,9 along
. deciduousforest fldec Sa.dasAlmas,Crateús,CE This study 14,6 2,6 2,2 22,5 58,1 a
2011 caatinga caa Faz.NãomeDeixes,Quixadá,CE Costaetal.(2007) 42,9 2,3 12,8 15,8 26,3
cerradofechado cerfec Brasília.DF Ratter(1980) inBatalha& Martins(2002) 0,7 1,8 44,9 13,5 39,1 topographic
cerradoaberto cerab PARNAdas Emas,GO Batalha& Martins(2002) 3,7 2 49,9 12,8 31,6
cerradoaberto cerab LagoaSanta,MG Warming(1892)inBatalha&Martins(2002) 4,6 5,4 55,1 6,1 28,8
cerradoaberto cerab Mojiguaçu,SP Mantovani(1983) in Batalha& Martins(2002) 7,8 2,1 47 12,2 30,9 gradient
cerradofechado cerfec Pirassununga,SP Batalhaetal(1997)inBatalha&Martins(2002) 5,6 1,1 36,1 17,1 40,1
cerradofechado cerfec Sta.RitadoPassaQuatro,SP Batalha&Mantovani(2001) 6,7 0,8 30 17.2 45,3
&
inBatalha Martins(2002)
pluvial forest flpl AltodoPalmital,FozdoIguaçu,PR Cainetal.(1956) 0 3 11 6 80
pluvial forest flpl Caiobá.PR Cainetal.(1956) 0 3 3 7 87
pluvial forest flpl Mucambo,Belém,PA Cainetal.(1956) 0 0,9 2,8 0,9 95,4
temperate forest fl temp HortoBotânico.Pelotas.RS Cainetal.(1956) 5 5 16 4 70
cerradão cerradão AguasdeSta.Barbara,SP MeiraNetoetal. (2007) 0 0 4 1,3 94,7
cerradosensustrictu cerss ÁguasdeSta.Barbara,SP MeiraNetoetal. (2007) 0 2,8 10,7 9,6 77
campocerrado cpcer ÁguasdeSta.Barbara,SP MeiraNetoetal.(2007) 0 6,4 19,2 14,1 60.3
camposujo cpsj ÁguasdeSta.Barbara,SP MeiraNetoetal.(2007) 0 7.9 31,8 41,3 19,1
campolimpo cllp ÁguasdeSta.Barbara.SP MeiraNetoetal.(2007) 5 0 32 34 14
restinga res Itamaracá.PE AlmeidaJRetal.(2007) 16,8 5,3 8 19,5 50,4
inselbergvegetation inselb Quixadá.CE Araújoetal.(2008) 44,2 2,6 13 15.6 24,7
cerradosensustrictu cerss Itirapina, SP Batalha& Martins(2004) 1,8 1.8 18,6 11.5 66,4
caatinga caa Betãnia.PE Costaetal. (2009) 40,5 1,1 14.6 18 25.8
restinga res Caravela.BA MeiraNetoetal. (2005) 9 0 14,9 23,9 52.2
restinga res Mucurí. BA MeiraNetoetal. (2005) 7,5 0 28.3 24,5 39,6
Note:cüannga-xencshrubiand.carrasco—deoduousshrubLmd:cerradosensustriclo—sa\anna:cerrado<fechado=densesa\anna;cerradoaberto=opensavanna;campocerrado=grasslandwilhscateredshrubs;camposujo=grasslandwilh ^
scateredshrubs:campolimpo=grassland:cerradão=tallwoodlandsa\anna:restinga=sandvCoastalplains.
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Araújo. F.S.etal.
Th Ci H Ch O Ph
CA CR FD CA/CR CATO CR/FD ALL
Occurancyclasses o 20 40% 60 80 100
species
Figure3 - Proportionoffamilies(white) and species Figure4-Life-formspectraofthethreephysiognomies
((bClAac-k)caoactciunrgraingCRin-ocnaer,rtaswcoo,orDiFn-aldlepchiydsuioougsnfoomreisets) (CA-caatinga CR-carrasco,DF-deciduousforest)of
ofSerradasAlmasNatural Reserve, Crateús, State of SerradasAlmasNaturalReserve,Crateús,StateofCeará,
Brazil, compared to Raunkiaer’s normal spectrum (N).
Ceará, Brazil.
Valuesovereachphysiognomybarindicatethenumber
ofspeciesofeachlife-form.Speciespercentagesofeach
woodyInstpheecipehsys(isohgrunbosmiaensdotnreseesd)ipmreendtoamriynraetlieedf,, tlihfeer-ofpohrymtaere(Tehx)p,rcersyspetdobpyhythteew(iCdr)t,hhoefmtihcerbyapr.toLpihfey-tfeor(mHs):,
totaling 124 in the deciduous forest and 68 in the chamaephyte(Ch),phanerophyte(Ph).
carrasco. In the caatinga, on the crystalline
basement, the highest species richness (69) was
represented by herbs. group, carrasco and deciduous forest exhibited
The life-form spectra of the studied scoresclosetothoseofrestingaanddifferentfrom
physiognomiesdifferedsignificantlyfromthenormal those ofcerrado, apparently because ofthe lower
spectrum (caatinga %2= 159.33 p < 0.01 df= 4; proportion ofcryptophytes (Fig. 5). The caatinga
carrascoX:=49.07p<0.01 df=4;deciduousforest composedawell-definedgroup,whichcomprised
X"= 120, p<0.01 dl =4). In general, thecarrasco spectra of other caatinga studies, including
andthedeciduousforestexhibitedsimilarproportions vegetation on inselbergs. This group is associated
of species ofeach life-form, whereas the caatinga with higher proportion of chamaephytes and
exhibitedadifferentspectrum(Fig.4).Therophytes, therophytes(Fig.5).
hemicryptophytes and chamaephytes were the
predominantlife-formsinthecaatinga(69%),carrasco Discussion
(53%)anddeciduousforest(46%),respcctively;thus, In general, in the semi-arid region of
they characterizeeach physiognomy.
northeastemBrazil,areaswithhigherannualaverage
Inthecomparisonsoflife-formspectraamong
rainiall associated with higher altitudes exhibit
pawxihetyshsioootfghtnehroemBiDreaCsziAolficaSonerrvrreeagsdepatosantAdielodnmattsoypoNevaset,urtrh6ael0t%RweoosfeFritrvhseet ahwli.ags2h0e0ra7lss;pFoeecriorebasszreeitrcavhl.ened1s9s9i8(n;LiGthmoeameeptshayl1.s982i00o)0.g9nT;hoAimrsiapeúasjtoteoemft
totncohfertexaylltispfeitencot-eoporfntohdiryat.m:thIs4eenp9set.och6atner8neao%dsrsdotiinooinfoita)dhtepiooouhfftniar:sndctiei)haarsaxgopimrepsacahatemynrptdathehw1rysi3et,t.ehe3g0ssiric%o)oaurnopoendssf adNsleaettcdiuiitrdmuaeudlnoetusRasertsfyheoarraenrvseetat.sah.Beneedsvccieaadnarctrswia,instcgdhoae,rcaiibinodnftuaSholeullrsoircnavadetegdexeadtesaasttAshiiliomgnmihlaoeasnrr
tothecaatingaarcaofthecrystallinebasement,have
tcoAhraldermirranoaaspsthciyNootanetasusnrpd(aaFlcitegh,R.eei5snd)e.gerrcTvoiehudepunoe2lua,irsfwlehy-fifoocroverhsmetarsllpisaneopcpctSeoredmarproiranfisdtteahhssee Tberixhecceehensnpeeptsoeisixoncn(tesSepid(tliRvoooaundtesatiisnahleg.otewna2le0.tr0ha3a1l)t9,9abs8et;ihhnaoPgvueirsgneehigdrtihahmieeegrntheteaaalr.rrsey2p0sae0lco2oim)nee.se
the restinga and cerrado spectra (Fig. 5). In this does not result in higher species richness; other
Rodnguésiá62 2 341-366 201
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Roristicsandlife-formsalonga topographicgradient
inwoodygrowth-formsisrelatedtotheincreasein
HHh +A OLtifhec-rfsotrumdies aridity (lowerrainfall and highertemperature). In
o SerradasAlma* previous studies, the replacemcnt of non-woody
ccrVfcc -cerab life-formsbywoodylife-formsandtheincrease in
erradio &“A cpAsjA cplp richness along humidily gradients have bcen
x observed inaridareas(Pavónetal.2000),tropical
< Ph \ lltemcppwr1^ |+Cr ^ inselb savannas (Williams et al. 1996), forests and
11P* cmt temperategrasslands(Kovács-Langetal.2000).
lldec O+C-li Considcring woody and herbaceous flora
together, the deciduous forest on lhe sedimentary
basin exhibited higherrichness than the caatinga
o.o 0.5 1.0 1.5 2.0 2.5 located on the crystalline basement. Potentially,
Axis 1 there must be higher humidity in the air and soil
Figure 5 - Ordination diagram of lhe detrended resulting from the elevation; there must be also
correspondenteanalysis(DCA),withscoresoflife-forms (solialtsowsiotlhspearnmdanqeunatrwlaztesraavnadisl)a,bilwihtyicinhdepeopsslaiybelrys
andcompiledinventorics,includingthephysiognomiesol
contributctothehigherfloristicrichnessobserved.
Serra das Almas Natural Reserve. Abbreviations tor
Comparing the carrasco and the deciduous
(veTghe)l,aticornytpytpoepshfyotlelow(CTra)b,leh1e.mLiifcer-yfportmosp:htyhteerop(hHy)t,e forest located in the same sedimentary basin, the
latter exhibited higher richness. In this case,
chamacphyte(Ch),phanerophytc(Ph).
humidity seems to be an important factor: lhe
deciduous forest is located on lhe cuesta and the
factorsmustalsobeconsidered,suchaslheposition carrasco on the immediate backside. On the
of the hogback, levei of desiccation of the relief backsidetheairisprobablydrierandwindspeedis
and physiochemical composition ofthe soil. The higher, which causes more desiccation. Besides,
dReesceirdvueouisslofcoraetsetdoonftSheerwrianddawsarAdlsmiades, bNeattwucreanl ssoainldiseere(mAsratoújpola&yaMarrotleintsoo1,9s9i9n;ceArcaaúrjroasectoals.oi1l9s99a)r.e
500 m and 700 m. whereas the carrasco, though Despite the high species richness found in
oloncattheedIatceawhairgdhesridaeltaitnuddeoanbosuatnd7i0e0r mso,iliss,lowchaitcehd thhieghrleiggihotn tohfeIbcioanptarbiabuPtliaotneauo,fitthies inmopnor-twaonotdtyo
arensdulstlseinndearpphlyasntisogannodmlyoowferlorwiechrnheesisghtth,ansmianllteher clioamnpaos)nctnotth(ehertbost,alssupbeschireusbsriacnhdnehsesrboafceeaocuhs
physiognomy. In lhe caatinga, on the crystalline
deciduous forest.
Conccmingtheherbaceouscomponcntoflhe basement. non-woody plants werc rcsponsible for
Braziliansemi-arid Hora,studiescarriedoutinthe most of the floristic richness, that is expectcd in
siinnetdneisrcu-aptlcsatttrehiaactutodtehipesrheiisngshiteohsnetohrfeitcrhhbenaeccrseyssotouafslltichneoemccpaoaomtnpiclnnegtxa oainrfittdhhacerncodpashreyrmtaiess-caoirniatdnhdcelsiienmcanttvhciesr,doedncumieendttuoso.tuhsOenpfortreehdsctoc.omniwtnoraoanrdcyye,
(Sampaio 995:Rodaietal.2005;Costaetal.2007: plantswerercsponsibleforthehighestrichness,since
Mamede &1 Araújo 2008). Comparatively, studies inmorehumidclimatesphanerophytcspredominatc.
craircrhineedssouotfinheserdbiamceenotuasryflfoorram(utRioodnasireetcoarl.de1d9l9o9w; cstablHiisghmhcenrt owfaltiefre-faovrmasiltahbaitldiotynotfanveoerdslatrhgee
reduetions of the aerial shool system during lhe
FigueiIrnedSoercrtaal.d2a0s00A)l.mas Natural Reserve, the unfavorable season (phanerophytcs), which is a
floristic richness of woody species incrcased at nccessarystrategy forlhesurvivalofmostspecies
high altitudes in areas of deciduous forest and inaridandsemi-aridregions(sceRaunkiaer 1934;
carrasco, whereas the richness of herbaceous van Rooyenetal. 1990;Kovács-Langetal.2000).
speciesdecrcased.Theincrcascinrichnessof trees In the case of Serra das Almas Natural Reserve,
and shrubs with altitude seems to bc a general whichisinsertedinasemi-aridclimaticdomain,the
patternforvegetationofaridandsemi-aridregions. increaseinaltitudemaypotentiallyfavorhighwater
In the Brazilian semi-arid region, the increase in availabilityonthewindwardside.Besides,soilmust
richnessofherbaceousgrowth-formsanddecrease betakenintoaccounl,sincetherearetwodiffcrent
Rodriguèsia62(2):341-366.201
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cm 12 13 14 15 16 17
1
348
Araújo. FS.etal.
geological units: lowlands of the crystalline deciduous forest occurs on the windward side
bascmeHnetrbaancdeothuesMoeriosuNbor-tweoosdedyimpelnatnatrsy(bhaesribns., dwhoenroctasftohremcadrirscarsectoeocunciutrss.oTnhtehyelfeoerwmaradcsiodnel),itnhueusme
subshrubsandherbaceouslianas)arethelife-forms represented by species overlap and by the same
sthyastteexhmibdiutrtihenghigthheestdrreyduscteiaosnoonft(hteheaerroipalhystheoso,t biologiWchalesnpeacntarluymz,iansgepmhpyhsasiiozgendombiyeAsusotnindi(f2f0e0r5e)n.t
c1r9y3p4t)o.phTyhteesb,ioalnodgichaelmiscpreycpttroupmhyotfest;heRacuanaktiiangear sgoeiolmocropmhpoolongeinctalmuaniyts,deaptaerrtmfirnoemdtihseccrleitmeatuen.itthse;
studied was characterized mainly by therophytes, communities that, according toWhittaker(1975),
a life-form characteristic of arid and semi-arid eanbedelimited by floristiccompositionand life-
regions (Raunkiaer 1934; van Rooyenetal. 1990; forms, such is the case of the difference found
Kovács-Langctal.2000). Indeed,amongthethree between the caatinga and the complex deciduous
physiognomics studied, the caatinga occurs on forest+carrasco.
shallow soils in the lowlands of the crystalline In the comparative analysis with the
basement,wheretemperature ispotentiallyhigher biological spectra from other Brazilian seasonal
andrainfall is potentially lowerthan in mountain- vegetationtypes,thediscriminationofthecaatinga
range areas, resulting in lower water availability. by higher proportion of therophytes and
The physiognomies on the Ibiapaba plateau chamaephytes shows that this vegetation is
(carrascoanddeciduousforest) mustoccurunder composed of species whose life-forms represem
lower water restrictions, since higher altitude better the semi-arid climatic pattern, since the
craoinntfrailbluteasndtoltohweeprotteentmipaelraotcucruer,renwcheicohf hfiagvhoerr pofrevdeogemtiantainocnesooffatrhiedseanldifsee-mfio-ramrsiidsecnhavriarcotnermiesnttisc
phanerophytes, a life-form characteristic of sites (Raunkiaer 1934; Cain 1950). The biological
withlowerwaterrestriction. spectrumissimilarlothespectrumofaridandsemi-
Inadditiontonumericdifferences in species aridclimatezonesoftheworld.
richness, remarkable differences between the Insummary,thetwogeomorphologicalunits
florislic complexes of each physiognomy were presentin thestudyareahavetwodistinct floristic
observed in the present study. The two main complexes, characterized by the predominanceof
complexes (caatinga and carrasco + deciduous therophytes on the crystalline basement and of
forest)areconsistemwiththesoil typesthatoccur phanerophytes on the sedimentary basin. These
in the area, resulting from the typeofsourcerock. results show that when implementing reserves in
Allhough species overlap between deciduous Braziliansemi-aridareas,abioticlocalfactors,such
forestandcarrascomaybeconsideredlow(15%), as soils and relief, must be taken into account,
differences are even larger when compared with because these lactors seem to reflect regional
caatinga,whoseoverlapisonly4%. Carrascoand floristicvariation.Theenvironmentalheterogeneity
deciduous forest are floristically more similar may result not only in high species diversity, but
becausebothhaveasetofspeciesthatprefersandy also in high functional diversity in the Brazilian
soil with low pH, whercas caatinga differs from semi-ariddomain,which, inthepresentstudy,may
that floristic group by the presence of species be observed in differences in life-form spectra
typical of soils originated from the crystalline among the three physiognomies analyzed.
basement of the inter-plateau depression. The
crystalline and sedimentary floras ofnortheastem Acknowledgements
oLABBlorbrrecaasamazzeúloiirjllsivoinea&veadntelnsRaistlonoe.odmar2daiini0-ieal0aslf52,rye0;isrc0deaL2sarri;treomiAgfaealiddcoebaonotrtfuaao(tolaA.rmidraa2neadt0srúor0ej-i9voFsc)eci.erealstalhlecao,lra.eertaae1staa9les.9di8to2faf0w,r0tao3hbsm;e; otrCheuaestaetarwrivieTneat,ghhaewtrnhaffsoruuonncnu-dadgrgierhnodigvewedthfmhoimreuecotnhm.mtLaatManhliteaenorigrf,sgelatosmnrétieiruszdnitaioittecisdopiewnlneAavCrsneiesnêotfcncooacirrriayraçtiãhoeeedof
BrazilAiasnAsnedmria-adrei-dLriemgaio(n1,9w81h)eenmtphheaspirzeeddo,miinntahme -T(eCcNnPolqo/gPiEa,LDlo-nPge-stqeurimseacEocloolgóigciaclardeeseLaornchgaprDougrraaçmãso
pvahryisaitoigonnoimsieisnsctluidmiaetde,inatsheoIbbsiearpvaebdaPilnattehaeu(ttwhoe C(NIsiMPtSeqEC(aAparRtoci-.nMgnCa°.)T,4/7IC6nN2st8Pi5qt/u)2t0,o0d3oo-f8M)EidlaiêntdnailPoRUdOnoiBvSIeerOmsi/aáMlriMddAoo
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JournalofBiogeography23:747-756.
Artigorecebidoem 09/05/2010. Aceitopara publicaçãoern 18/12/2010.
Rodriguèsia62(2):341-366.201
SciELO/JBRJ
cm 12 13 14 15 16 17