Physiognomy and structure of a seasonal deciduous forest on the Ibiapaba plateau, Ceará, Brazil / Fisionomia e estrutura de uma floresta estacional decídua no planalto da Ibiapaba, Ceará, Brasil

2011·4.3 MB·Portuguese

by  Jacira Rabelo Lima

)
Rodriguésia62(2):379-389.2011
httpI//rodriguesia.jbrj.gov.br
Physiognomy and structure of a seasonal deciduous forest
on the Ibiapaba plateau, Ceará, Brazil1
FisionomiaeestruturadeumaflorestaestacionaidecíduanoplanaltodaIbiapaba,Ceará,Brasil
JaciraRabeloLimaz Everardo ValadaresdeSá BarrettoSampaio ,
MariaJesusNogueir,aRodai2&FranciscaSoaresAraújo
Abstract
TheBrazilianscmiaridregionisdominatedbycaatinga.However,othervegctationformationsoccur,induding
deciduous and semi-deciduous forests. This study describes physiognomy and structure ofa forest on the
sedimentary Ibiapabaplateau. All plants within one hectare were separatcd into thrce componcnls: woody
plantswithperüneteratsoillevei(PSL)>9cm(WCLP),woodyplantswithPSL>3and<8.9cm(WCSP),
andhcrb/subwoodyplants(HSwC).WCLPincluded88species(33families),WCSP50species(23families)
andHSwConly7species(5families).Totaldensity,basalarea,andmaximumandaverageheightofWCLP
were5683 plants/ha,47 mVha, 18and 5 mrespcctively.TotaldensityandbasalareaofWCSPwere 17500
plants/haand2.8rnVha,respectively.DensityofHSwCwas9plants/m2andonly31%ofthesampledarcawas
occupied by this component.
Keywords: basalarea;drytropical forest;phytosociology;plantheight.
Resumo
Osemi-áridobrasileiroédominadopelacaatinga.Entretanto,outrasformaçõesvegetacionaissãoencontradas,
porexemplo asflorestasdecíduasesemidecíduas.Esteestudodescreveafisionomiaeaestruturadeumafloresta
estacionainòplanaltosedimentardaIbiapaba,Ceará.Foiselecionadaumaparceladeumhectareeasplantas
separadas em três componentes: plantas lenhosas com perímetro ao nível do solo > 9 cm (CLS), plantas
lenhosascomperímetro>3e<8.9cm(CLI),ehcrbáccas/sublenhosascomperímetro<3cm(HSL).NoCLS
effao5mríalmimase)rn.ecsoNpnoetcrtCaiLdvaSas,me8dn8etnees.sipdNéacodieeCsLto(It3,a3l,dfeáanrmíselaiidabasad)se,alneoeáaCrleLtaIurb5aa0smaáelsxpiféomcriaaesmc(m12é73d5if0aa0mífploliraaansmt)ae5s/6nh8oa3HepSl2a,Ln8tsamest/eh/haea,s.p4éE7cimmesH/(ShcaLi,,n1c8oa
densidade foi9plantas/m2eapenas31%daáreaamostrada foicobertaporessecomponente.
Palavras-chave;áreabasal; florestatropicalseca; fitossociologia;alturadeplanta.
belong to two physiognomic types: 1) non-forest
Introduction
formations, mainly savanna (cerrado and closed
The heterogeneity of the flora and shrubland(carrasco),onthesedimentaryplateaus;
pinhytsheioBgrnaozmiliieasnosfemtihaerididffreergeinotnviesgceatautsieodnsbytytpweos oannds2e)dipmeernentnairayl,ansedascroynsatlalfloirneestsufbosrtmraattieosns(,Robdoatih
rainfall gradients, one in the South-North and the &Nascimento2002;Araújoetal. 2005b).
other in the East-West direction, and by tnarked Flora, physiognomy, and structure of these
geologicdifferences(Rodaietul.2008).Athighcr forestsarescarcelyknown(Andrade&Rodai2004;
altitudes, where aridity is less accentuated, the Rodai & Nascimento 2006), espccially those
u2sl0e.a0s21o;9n9Fa8el;rrnAarozaneú-tjtuohl.oetm2y0ul0.3f;o1r9Am9ra9at;úijRoonosdeatoiucl.c&u2r0N0a(5sAbcr)ia.úmTjehoneteyot 2oc0c0u4r)r.inMgoorneosvedeirm,entthaerrycpilsatneoauspu(bAlnidsrhaedded&etRaoidlaeid
1PUanritvoefrtshicdMaadestFcerdetrheas]isRourfatlhcdcftrPsteraunlahmobruactoU,niPvceprtsoi.daBidoelFoegdiear,aRl.RDuroalmdMeaPneoren,amdbeuMceod.eiross/n.Doisirmãos,5«2n1i71o9o0o0,RReecciiffee,PPEE,BBrraassiill.
'UniversidadeFederaidePernambuco,Depto.EnergiaNuclear,Av.Prof.LuisFreire1000,50740-540,Recife.PE,Brasil.
*UniversidadeFederaldoCeará,Depto.Biologia,Av.MisterHulls/n.60455-760.Fortaleza.CE.Brazd.
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Lima.J.R. etal.
description of the vertical organization in these Thesoi]wascharacterizedbydigginga1 xI m
forests whichhelpdesign projects to manage these trench and collecting samples from the top 10-cm
forestsinordertomaintainbiologicaldiversity. layerand from the layerbelowdown totheparent
The description and classification of plant material (75 cm depth). Physical (Tab. 1) and
communities generally focuses on features such as Chemical (Tab. 2) analyses were performed in the
floristic composition, structure and relative species Departamento de Ciências do Solo. Universidade
abundances(Box&Fujiwara2005).Thespecieshave Federal do Ceará. The soil was classified as a
different positions along a vertical gradient oflight dystrophicLatosol,poorlydeveloped,withlowpH
intensity,occurringonehigherthanlheothertoform and low cation exchangecapacity (T). In general.
thecommunity’sverticalstructure(Whittaker1975). Ca, Mg, K, P,organic matterand nitrogen content
The vertical differentiation is most pronounced in werelowanddecreasedfromthesuperficial tothe
woody vegetation that has various synusia which subsuperficial layer, while the opposite occurred
mayhave floristiccompositions independemofone withAlcontent.Texturevariedfromsandytosandy
another (Maarel 2005). Based on the vertical loam. Coarse sand, silt andclaycontent increased
stratificationofthe vegetation, it is possible to infer with depth, while that ol fine sand decreased.
the potential composition of functional groups of
differentstrata.Thevegetationcanthenbemanaged
Phytosociological survey
dainenocdridsdteuTrrohutucuostsmu,faroietrnheotissfatiwtonhofrmrteakheexadiIiibfmmfisuearmpteaonlbdtoacesasstlcerbrdaiitiobamleoeognfpithtcahyarelsyidsipeovlageasrntsoeoinatmauyyl.. aw)hwicohoTdchyoemcppolraminptsosendweeanrltel pwdliiavtnihtdlseadwrgiinetthpoestrhitremeemetcpeoermr(piWomCneetLnePtr)sa,:t
CWfeohraaerstát.?aWr2e)etIhaseitmphhitsyosfaoinrosegswtneorsmttryhuecatfnuodrlaltlohlweyissntigrmuqiculteausrrtetiooonofst:thhei1rs cswoooimlopdrlyeivseciinogm(pPpSolaLnn)etnsetwqiuwtaihlthPtoSsLmoar>lgl3repaaentrdeir<m8ett.he9arnc(m9W;cCamSn:Pd)bc,))
herb/subwoody component (HSwC), comprising
forestformationsoftheBraziliansemiariddomainor
tonon-forestformationsofothersedimentaryareas? plants with green stcms, without or with slight
lignificationintheaerialpartandthatwereupto 1 m
Material and Methods talland2.9cmperimeter,cxcludinglheyoungplants
ofwoody species.
Study area The study hectare was divided into plots
Thestudy areais located within the 2794ha ofdifferent sizes, depending on the component
odfasseAalsmoansa,lCfoeraersátSitnastied,eBtrhaeziRle.sTehrevaseNaastounraallfSoerrersat a1n0alxyz1e0d.mWplCoLtsPawnadsWanCaSlyPzeidn i5n01p0l0otcso,n2tixgu2oums
occupics a narrow strip in the upper part of the each,placed in the right proximal cornerofeach
steep castem slope on the south-central Ibiapaba alternate larger plot. In these two components,
plateau, which forms the oriental border of the theheightand PSLol all liveplants(exceptlianas)
Middle Northern Sedimentary Basin (Lins 1978). were measured, followingthecriteriapreviously
Tvehgeetaftoiroens,toins tloipmiotfedthetopltahteeauw,esatndbbyyccaarartaisncgoa d1emscreiabcchd,.aHtSthweClewftapsraonxailmyazledcoirnne1r00ofpleoatcsh1oxf
vegetation to the east, on the lower parts of the the larger plots, during the rainy season. In
slope, extending tocover most ofthe lowlands in HSwC, all plantswere identifiedandtheplotarea
Ceará. In the forest area, a one-hectare plot was proportion covered by each species was
installed, at an altitudeofabout 650m, within the estimated, withthehelpofa 1 x 1 mgrid,divided
following coordinates 40°54’5’’W and 5°8'29”S; into 100 squaresof0.1 xO.l m. Presence in one
40°54’45”W and 5°8’30”S; 40°54,46”W and ofthe squares w'ascounted as %coverage.The
1
5°8’36”Sand40°54’50"Wand5°8’35"S. botanicalmaterial wasincorporated intotheEAC
Mean annual rainfall in the study area from (Prisco Bezerra)herbarium,ofthe Universidade
2000to2004was 1044mm,JanuarytoAprilbeing Federal do Ceará. APG III (2009) classification
therainiestperiod,correspondingtomorethan80% system was adopted.
of annual precipitation. In general, rainfall was Thefollowingphytosociological parameters
concentrated in a single month and did not occur were calculated for WCLP and WCSP: relative
fromJulytoDecember.Themeanannualtemperature density (ReD, %), relative frequency (ReF, %).
duringthestudyyear(2004)was24.8°C. relative basal area (ReBa, %) and importance
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Physiognomyandstwcture ofaseasonaldedduous forestin Ceará. Brazil *M1
Table 1-ParticlesizeanalysisofsoilsamplesfromtheforestseasonaldeciduousmontancforestofReservaNatural
Serra das Almas, Crateús, Ceará State.
Depth(cm) Coarsesand (g/kg) Fine sand (g/kg) Silt (g/kg) Clay (g/kg) Texture class
Oto 10 130 710 80 80 Sand
11 to 75 240 510 140 110 Loamysand
Table2-ChemicalanalysisofsoilsamplesfromtheseasonaldeciduousmontaneforestofReservaNaturalSerradas
Almas, Crateús, Ceará State.
Depth Sorptive Complex p v M C N pH
(cm) Ca2* Mg2* Na* K* Al3* (mg/kg) (%) (%) (cmol.kg )
(cmol.kg')
0to 10 0.9 0.9 0.1 0.09 0.85 3 32 30 10.6 1.1 4 6.3
27 46 6.48 0.7 4.2 6.4
11 to75 0.7 0.9 0.1 0.05 1.45 I
Ca’’ =calcium; Mg2*=magnesium;Na -sodium; K potassium, Al"=aluminum; P=phosphorus; V=basesaturation;
M = aluminum saturation; C =carbon; N =nitrogcn; pH = soilpH; T = cation exchange capacity.
value (IV, %), using the formulas described by Piptadenia moniliformis, which accounted for
Rodai etal. (1992), and Shannondiversityindex, 28% of the total IV (Tab. 3). Gymnanthes sp.l,
according to Magurran (1988). The calculations Bauhiniapulchella and Crotonargyrophylloides
weredoneusingFITOPAC(Shcpherd2006). were the species with the highest absolute
AimingtocompareWCLPphysiognomyand frequencies(99%;97%and84%,respectively).m
strueture of the study area with that of other Maximumandaverageheightwere 18and5
seasonal forests and non-forest formations within (+ 2), respectively, and only 1 1% of the plants
the Brazilian semiarid region (only surveys with attained heights over8 m. Maximum and average
cinocnltuasiionnicnrgiteirnifonoorfmaPtSiLo>n3ocnm)toattaalblpelwaanstodregnasniitzye,d pdliaanmtestebreslwoenrgien6g5.t2oatnhde83.4tcom6(c6m),wdiilahme3t3e%r colfaaslsl
community basal area, maximum plant height, and47%tothe6to9and9to 12cmclasses. Inthe
proportion ofplants over 8 m tall. and mean and sameWCLPcomponent,twostratawereidentified.
maximumstemdiameters.Foreachcomparedsite, The lowest stratum was dominated by plants at
information on altitude, mean annual rainfall and most 8 m in height, with a continuous canopy
sample area were also included. Ten areas were distribution. The most abundant species in this
included in the comparison, five classified as stratumwereGymnanthessp.I,Bauhiniapulchella,
seasonalforestsandFiveasnon-forest formations. Croton argyrophyloides and Maytenus sp. Thme
upperstratumwasdominatedbyplantsupto 12
Results tall but some plants of Brosimum gaudichaudii,
Piptadenia moniliformis and SwartztaJlaenungii
(faWmCiLliPIe)ns,t(hT8ea8wbo.spo3ed)c.yieTcshoewmepfroaenmeifnloituenwsdi,rtihbcehlelasrogtnegipinensrgpiemtceoit3eesr ASwsweparrietdztoaislapleferlr,amweaimstiuhnobguiitnicfaaonnrdummTi,hnPigilpaotacaodgneltnaiiuancumooocunaisrlcpiaafnowroempirys.e,
wHerryet:hrFoaxbyalcaecaeeae(1a9nsdpeMcyirest)a,cEeuapeho(sribxiaeacceha)e.(T10h)e, the moTshtefwreoqoudeyntcsopmepcoienseninttwhiitshuspmpaelrlspterraitmume.ter
Shannondiversity index(H’)was3.20nats/plant. (WCSP) included 50 species, belonging to 22
aTontdal47denmsVihtay,anrdesbpaescatlivaerleya.weSrpeec5i6e8s3wpiltahntt/hhea families. The families richest in species were:
GhiygmhneasntthIVesanspd.lr,elBaatuihveinbiaasaplulacrheealslawearnde EFraybtahcreoaxeylaancdeaEcup(hfoorubr)iaacnedaeBi(gcingohntisapceccaiees(teharcehc)),.
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Tablc3-Phytosociologicalparametersofplantswithstemdiameters>9cminlheforestofReservaNaturalSerradas
Almas,Crateús,CearáState,indecreasingorderoftheirimportancevalue(IV).N-numberofindividuaisperhectare;
NP-numberofplotswherethespecieswasfound; ReD-relativedensityofthe species(%); ReBa-relativebasal
arcaofthespecies(%);andReF-relativefrequencyofthespecies(%).Collectors: FSA-FranciscaSoaresAraújo;
JRL-Jacira Rabelo Lima; LWLV -Luis Wilson Lima-Verde; MSS-Melissa S. Sobrinho; and SFV- Sandra
Freitas Vasconcelos.
N" Species/ Family Voucher IV N NP ReD ReBa ReF
1 Gymnanthessp.1 (Euphorbiaceae) JRL29 50 1507 99 26.52 18.2 4.88
2 BauhiniapulchellaBenth.(Fabaceae) JRL45 19 537 97 9.45 4.45 4.78
3 PiptadenianumiliformisBenth.(Fabaceae) FSA1298 17 122 70 2.15 11.1 3.45
4 CrotonargymphylloidesMüll.Arg.(Euphorbiaceae) FSA1294 15 333 84 5.86 4.94 4.14
5 Maytemssp.(Celastraeeae) JRL 100 12 279 76 4.91 3.31 3.75
6 Thiloaglaucocarpa(Mart.)Eichler(Combretaceae) LWLV1050 11 138 57 2.43 5.36 2.81
7 Erythroxyliunef.vacciniifoliumMart.(Erythroxylaceae) JRL69 9.9 167 73 2.94 3.37 3.6
8 ArrabidaeadisparBureauexK.Schum.(Bignoniceae) JRL20 9.6 203 80 3.57 2.1 3.94
9 AspidospennadiscolorA.DC.(Apocynaceae JRL 18 8.9 139 55 2.45 3.72 2.71
10 Eugenia cf.piauhiensis0.Berg(Myrtaceac) JRL62 8.1 158 72 2.78 1.79 3.55
11 SwartdaflaemingiiRaddi(Fabaceae MSS262 7.9 99 58 1.74 3.31 2.86
12 Xylosmaciliatifolia(Cios)Eichler(Salicaceae JRL77 7.3 189 65 3.33 0.8 3.2
13 CopaiferamaniiHayne (Fabaceae) JRL38 7.3 79 51 1.39 3.41 2.51
14 Buchenaviacapitula(Vahl)Eichler(Combretaceae) MSS292 7 31 28 0.55 5.04 1.38
15 Eugeniaaff.malhaCambess.(Myrtaceae) JRL73 6.7 133 60 2.34 1.36 2.96
16 AlibertiamyrciifoliaSpruceexKSchum.(Rubiaeeae) JRL 102 63 115 63 2.02 1.36 3.1
17 Eugeniaalf.dysentericaDC.(Myrtaceae FSA1291 6.5 94 43 1.65 2.71 2.12
18 Aspidospennasuhincanum Mart. (Apocynaceae) JRL 17 6.1 74 44 1.3 2.62 2.17
19 CombretumleprosumMart.(Combretaceae) JRL74 5.1 101 51 1.78 0.78 2.51
20 EphedrcmthuspisocarpusR.E.Fr.(Annonaceae) JRL 16 4.6 57 41 1 1.6 2.02
21 AcacialangsdorfiiBenth.(Fabaceae) JRL40 4.6 74 26 1.3 2.01 1.28
22 BrosimumgaudichaudiiTrécul.(Moraccae) Probio306 4.2 40 33 0.7 1.84 1.63
23 PeixotoajussieuanaMart.exA.Juss.(Malpighiaceae) JRL33 3.9 74 42 1.3 0.56 2.07
24 CrotonnepetifoliusBaill.(Euphorbiaceae JRL28 3.8 75 39 1.32 0.57 1.92
25 DalbergiacearensisDucke(Fabaceae) LWLV1070 3.6 45 34 0.79 1.1 1.68
26 ByrsonimaganlnerianaA.Juss.(Malpighiaceae) JRL32 32 43 36 0.76 0.66 1.77
27 HymenaeaeriogyneBenth.(Fabaceae) JRL42 3.1 42 22 0.74 1.3 1.08
28 EugeniaaurataO.Berg(Myrtaceae) JRL60 3 46 34 0.81 0.48 1.68
29 AgonandrabrasiliensisMiers(Opiliaceae) JRL56 2.8 31 21 0.55 1.23 1.03
30 VitexschauerianaMoldenke(Lamiaceae) JRL64 2.7 32 22 0.56 1.08 1.08
31 AspidospennamultiflorumA.DC.(Apocynaceae) JRL 19 2.6 19 12 0.33 1.63 0.59
32 Galipeaalf.trifoliataAuble.(Rutaceae) JRL36 2.3 30 21 0.53 0.72 1.03
33 Guapiragracilijlora(Schmidt)Lundell(Nyctaginaceae) JRL34 23 31 25 0.55 0.5 1.23
34 ErythroxyliunstipulosuinPlowman(Erythroxylaceae JRL86 2.3 45 26 0.79 0.2 1.28
35 ManihotpalmataMull.Arg.(Euphorbiaceae) JRL26 2.2 33 29 0.58 0.22 1.43
36 Ipoinoeabrasiliana(C.Martius)Meisner(Convolvulaccae) JRL25 2 37 24 0.65 0.18 1.18
37 Secorulontiací.foliosaA.DC.(Apocynaceae) JRL89 1.8 36 21 0.63 0.17 1.03
38 Dalbergiasp.(Fabaceae) JRL41 1.6 23 16 0.4 0.45 0.79
39 Bauhiniasp.(Fabaceae JRL44 13 46 10 0.81 0.23 0.49
40 Arrabidaeachica(Humb.&Bonpl.)B.Verl.(Bignoniceae JRL21 13 25 19 0.44 0.11 0.94
41 MachaeriumacutifohumVogei.(Fabaceae) JRL80 13 20 16 0.35 0.33 0.79
42 Tocoyenaformosa(Chain.&Schltdl.)K.Schum.(Rubiaeeae)JRL59 1.3 17 14 03 032 0.69
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Physiognomyandstwctureoía seasonaldeclduous forestin Ceará. Brazil
Voucher IV N NP RcD ReBa RcF
N° Spccies/Family
43 PaulliniacearensisSomner&Ferrucci(Sapindaeeae) JRL34 1.2 20 14 0.35 0.19 0.69
44 Eugeniapunicifolia(Kunth)DC.(Myrtaceac) JJRRLL6237 01.9 1166 1113 00..2288 00..112 00..6544
45 Gyrrmanthessp.2(Euphorbiaceae)
MSS248 0.8 13 II 0.23 0.07 0.54
46 TrigonianíveaCambess.(Trigoniaceae)
JRL79 0.8 13 10 0.23 0.11 0.49
47 CrotongrewioidesBaill.(Euphorbiaceae)
48 TassadiabnreheliiE.Foum.(Apocynaceae) JRL13 0.8 9 8 0.16 0.26 0.39
JRL65 0.7 12 10 0.21 0.03 0.49
49 TumerablanchetianaUrb.(Tumeraceae)
JRL87 0.7 11 9 0.19 0.07 0.44
50 Erythroxylumsp.(Erythroxylaceae)
51 Cnidoscolusvitifolius(Mill.)Pohl(Euphorbiaceae) JFRSLA514309 00..77 710 710 00..1182 00..2034 00..4394
52 Ourateasp.(Nyctaginaceae)
JRL51 0.7 12 8 0.21 0.05 0.39
53 HelicteresheptandraL.B.Sm.(Malvaceae)
JRL55 0.6 12 6 0.21 0.11 0.3
54 XimeniaamericanaL.(Olacaceae)
JRL37 0.6 8 8 0.14 0.08 0.39
55 DiocleamegacarpaRolfe(Fabaceae)
56 Jacarandajasnúnoides(Thunb.)Sandwith(Bignoniceae) JRL22 0.6 8 8 0.14 0.03 0.39
57 Justiciastrobilacea(Nees)Lindau(Acanthaceae) JJRRLL5135 00..55 75 75 00..1029 00..0124 00..2354
5589 AOmurbautreaancaf.cpeaarrveinfsoilsiaAElnlgeLmã(Noy)cAt.agCi.nSacme.a(eF)abaceae) JRL72 0.5 5 5 0.09 0.14 0.25
JRL61 0.5 6 6 0.11 0.07 0.3
60 Campomanesia sp. (Myrtaceae)
JRL35 0.5 6 6 0.11 0.05 0.3
61 Paulliniacf.elegansCambess.(Sapindaeeae) JRL43 0.4 6 5 0.11 0.06 0.25
62 HymenaeavelutinaDucke(Fabaceae) JRL58 0.4 5 5 0.09 0.02 0.25
63 Gouaniasp.(Rhaninaceae JRL78 0.4 4 4 0.07 0.09 0.2
64 KollinialeptopetalaR.E.Fr.(Annonaceae) JRL68 0.3 3 3 0.05 0.09 0.15
65 ErythroxylumbezerraePlowman(Erythroxylaceae)
JRL 104 0.3 5 3 0.09 0.05 0.15
66 Stachyarrhenacf.spicotaHook.F.(Rubiaceae) JRL52 0.3 4 4 0.07 0.02 0.2
67 HelicteresmuscosaMart.(Malvaceae) JRL66 0.3 4 4 0.07 0.01 0.2
68 ErythroxylumlaetevirensO.E.Sehulz(Erythroxylaceae) JRL 107 0.3 4 3 0.07 0.05 0.15
69 Arrabidaeasp.(Bignoniceae) JRL49 0.2 2 2 0.04 0.11 0.1
70 LonchocarpusararipensisBenth.(Fabaceae) JRL46 0.2 3 3 0.05 0.01 0.15
71 SertnacearensisAfran.Fem.(Fabaceae) JRL31 0.2 3 3 0.05 0.01 0.15
72 TrichiliaelegansA.Juss.(Meliaceae) JRL 103 0.2 4 2 0.07 0.04 0.1
77773564 SCCRuaoobpcmiihmaulicmoepsahaepfoTe.rraamrulgemuptvtiuotmpihf(loMloüieluolms.((WAMiralgrl.dt)..))HSuJp.brBe.ernGgi(.lEl(ueBtpithxo(arBebueiraasecc)reaacee)ae) JSJJRRRFLLLV4784852 0000....1222 4222 221i 0000....00004744 0000....1000322 0000....110055
777978 LCTuraeobttezobeunlibbaulrcafgn.icoahceahtuririaacncuuelsaaM(tüCalh(laA.ilnAl.re)gm.Sãt(oaE)nudDplhu.oc(rkBbieiga(ncFoeanbaiaecc)eeaaee)) JJRRLL2713 00..11 11 11 000...000222 000...000735 000...000555
JRL76 0.1 1 1
80 Lindackeriaovata(Benth.)Gilg (Achariaccae) JRL39 0.1 1 1 0.02 0.02 0.05
81 Mimosasp.(Fabaceae) MSS248 0.1 1 1 0.02 0.01 0.05
82 ColubrinacordifoliaReissek(Rhamnaceae) 0.02 0.01 0.05
JRL24 0.1 1 1
83 CondiarufescensA.DC.(Boraginaceae) 0.02 0 0.05
84 Peltogyneconfertiflora(Mart.exHayne)Benth.(Fabaceae) JRL50 0.1 1 1 0.02 0 0.05
85 ErythroxylumbarbatumO.E.Sehulz(Erythroxylaceae) JRL67 0.1 1 1 0.02 0 0.05
&Bamcby JRL47 0.1 1 1
86 Sennatrachypus(Mart.exBenth.)H.S.Irwin
(Fabaceae) 0.02 0 0.05
JRL57 0.1 1 1
87 BredemeyeraijloribundaWilld.(Polygalaceae) 0.02 0 0.05
JRL81 0.1 1 1
88 CrotonbetaceusBaill.(Euphorbiaceae)
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TheShannondiversityindcx(H')was3.26nats/plant. comparedarcas,wrasfoundinSerradasAlmasbut
Total densily and basal area were 17500 plant/ha theproportionoftreesover8mtallwaslowerthan
and2.8nr/ha,respcctively.Maximumandaverage thatoftheotherforests,except SerraTalhada,but
heights were 5 m and 1.4 m (± 0.7), respectively, higherthan that ofthe non-forest formations.
mostoftheplantsbcingunderthan2 m tall (83%) In Serra das Almas two distinct strata are
andlessthan 1.6cmdiameter(70%).Specieswith distinguished,ashasbecnregisteredforNeotropical
the highest IV were Lanterna sp., Xylosma deciduousforestselsewhere(e.g.,Murphy&Lugo
ciliatifolium and Croton argyrophylloides, which 1986), that are essentially trec-dominated with a
accounted for 32% of the total IV (Tab. 4). more-or-less continuous canopy and in which
Cranocarpus gracilis, Lantana cf. brasiliensis, grassesareaminorelement(Mooney etal. 1995).
Lantana sp„Justiciafragilis and Wedelia villosa In non-forest formations in Northeast Brazil.
wereexclusivetothiscomponent. especially closed shrubland (carrasco), only one
The herb/subwoody component (HSwC) stratum is recognized. The presence of scattered
had only seven species, belonging to 5 families very tall trees emerging above the upper stratum
(Tab. 5), excluding saplings and scedlings of distinguishes Serra das Almas from other
speciesthatreachlargersizes.ThefamiliesPoaceae deciduous forests in the region.
and Bromeliaceae had higher species richness for The woody vines formed a considerable
WCLP
this component (two species each). The Shannon proportionofthe species,with 13 species
diversity index (H’) was 1.16 nats/plant. Density (16%),distributed in ninefamilies,corroborating
was9 plants/m: and only 33% ofthe sample area Gentry‘sstatement(1982, 1995)thatvinesarean
was occupied by this component. Most of the important component in Neotropical seasonal
forests, where they represem about 20% of the
plants (63%) belonged to three species
species.
(Streptostachys asperifolia, Pavonia sp. and Some of the highest IV species, in the
Scaphispatha hastifolia), which, together with WCLP, in Serra das Almas (Arrabidaea díspar,
Bromelia auriculata, accounted for most of the
Croton argyrophylloides, Piptadenia
plant coverage ofthis component.
moniliformis, Rollinia leptopetala and Thiloa
glaucocarpa are also important species in
Discussion
Northeastern non-forest formations (Oliveira et
SerradasAlmasforestisphysiognomically al. 1997; Araújoetal. 1998;Araújoetal. 1999),
distinct from other seasonal montane forests and mainly those on the Middle Northern
also from all non-forest formations already Sedimentary Basin,whichextendsfora largearea
describedinNortheast Brazil. Ingeneral,seasonal west of the Serra das Almas location. Other
montane forests are physiognomically more important species, like Aspidosperma discolor,
variablethanrainforests,varyingfromtall forests A. subincanum and Brosimum gaudichaudii,
to low scrub(Pennington etal. 2009). Density of although present in these non-forest formations,
the most conspicuous component. trees and occurwith much lowerdensities.
shrubswithlargeperimeters(WCLP),washigher On the other hand, many of the important
than those of the other forests (Tab. 6), exccpt speciespresentintheseformationswerenot found
onelocatedat 1 lOOminacrystallinemountainin in the Serra das Almas forest. In spite of these
Serra Talhada, Pernambuco (Ferraz etal. 2003), differences,thefloraoftheSerradasAlmasforest
but lower than those of non-forest formations, is relatively similar to that of the non-forest
especiallycarrasco(Araújoetal. 1998;Araújo& formationswhileitisverydifferentfromthatofall
Martins 1999). other deciduous montane forests studied. with
Basal area, on the contrary, was lower than which it shares only a few species, none ofthem
thatofotherforests,exceptforbasal areaat900m withhigh IVeitherin SerradasAlmasorin those
inSerraTalhada,buthigherthanthatofnon-forest forests(Limaetal. 2009).
formations.Consideringthatbasalarcaisaproduet Comparisonsoftheothertwoplantcomponcnts.
of density and stem diameter, the average stem thewoodycomponentwithsmallperimeter(WCSP)
diameterinSerradasAlmasissmallerthanthatof and theherb/subwoodycomponent (HSwC). with
the other forests but greater than that ofthe non- thoseofotherNortheasternformationsaredifficultto
forestformations.Thetallestregisteredtree, in all makebecauscofthescarcityofthesemeasurements.
Rodriguésla62(2):379-389.201
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1 ) )
Physiognomyandstnicture ofa seasonaldedduous forest in Ceará, Brazil
NTaatbulreal4S-erPrhaytdoassocAilomlaosg,icaCrlaptaeúrsa,meCteearrsáoSftaptlea,ntisnwdietchresatseimngdioarmdectreorfst>he3ircimmpaonrdta<nc9ecvmalmuet(hIeV).orNest-onfuRmebseerrvoaf
FiRnredaBinvaciids-ucaraeilsSaoptaiervreehsbeacAstraaarlúej;aor;NeaPJRo-Lfntu-hmJeabcseiprreacoiRfeapslboe(tl%so)w;Lhiaemnrade;RtLheeWFsLp-eVrcei-leaLstuiwivasesWfifroleuqsnuoden;nLeRy.emoaDf--Vterheerldasetp;tevceMtdeSseSns-UMyeloC.fosltslhaeecSst.opreSseo:.benFsnS(h/A„o);;-
and SFV - Sandra Freitas Vasconcelos.
No Species/ Family ___ Voucher IV N NP ReD ReBa ReF
JRL 109 46.45 52 29 14.86 19.09 12.5
1 Lanhinasp.(Verbenaceae)
JRL77 24.86 30 15 8.57 9.82 6.47
2 Xylosmaciliatifolia(Cios)Eichler(Salicaceae)
FSA 1294 23.87 31 13 8.86 9.41 5.6
3 CmtonargyrophylloidesMidiArg.(Euphorbiaceae)
4 CranocarpusgracilisA.Fernandes&P.Bezerra(Fabaceae) JRL84 21.02 28 16 8 6.12 6.9
JRL29 19.13 28 13 8 5.53 5.6
5 Gymnanthessp.1 (Euphorbiaceae) JRL22 13.53 15 9 4.29 5.37 3.88
6 Justiciastmbilacea(Nees)Lindau(Acanthaceac)
JRL81 11.64 12 11 3.43 3.47 4.74
7 CmtonbetaceusBaill.(Euphorbiaceae)
JRL85 8.23 12 7 3.43 1.79 3.02
8 WedeliavillosaGardner.(Asteraceae)
9 AlibertiamyrciifoliaSpruceexKSchum.(Rubiaceae) JRL 102 7.75 8 8 2.29 2.01 3.45
JRL28 7.65 8 8 2.29 1.92 3.45
10 CrotonnepetifoliusBaill.(Euphorbiaceae)
JRL83 7.54 9 8 2.57 1.52 3.45
11 EvolvulusmacmblepharisMeisn.(Convolvulaceae) JRL45 7.24 8 8 2.29 1.51 3.45
12 BauhiniapulchellaBenth.(Fabaceae JRL20 6.32 7 6 2 1.73 2.59
13 ArrabidaeadisporBurcauexK.Schum.(Bignoniaceae) JRL51 5.85 6 6 1.71 1.55 2.59
14 HelictereshepumdraL.B.Sm.(Malvaceae) JRL25 5.42 8 1 2.29 2.7 0.43
15 JusticiafragilisWall.exClarke(Acanthaceac) JRL64 5.05 5 4 1.43 1.9 1.72
1167 BVainteixstsecrhiaoupesriisacnf.asMtoellldaerinsk(eGr(iLsaembi.)acBe.aGea)ttes(Malpighiaccac JJRRLL71301 44..8933 45 44 11..1434 11..7987 11..7722
18 Eugeniaaff.uvalhaCambess.(Myrtaceac) JRL 108 4.61 5 4 1.43 1.46 1.72
19 Lanternaaff.brasiliensisLink(Verbenaceae) JRL65 4.61 5 5 1.43 1.03 2.16
20 TumerablanchetianaUrban.(Tumeraceae)
JRL44 4.47 7 2 2 1.61 0.86
21 Bauhiniasp.(Fabaceae)
JRL 100 4.36 4 3 1.14 1.92 1.29
22 Maytenussp.(Celastrueeae)
JRL41 4 4 3 1.14 1.57 1.29
23 Dalbergiasp.(Fabaceae)
JRL34 3.88 3 3 0.86 1.72 1.29
24 Guapiragraciliflora(Schmidt)Lundell(Nyctaginaceae)
JRL42 3.63 4 4 1.14 0.76 1.72
25 Ipomoeabrasiliana(C.Martius)Meisner(Convolvulaceae)
JRL52 3.38 4 3 1.14 0.95 1.29
26 HelicteresmuscosaMart.(Malvaceae)
JRL33 3.11 3 3 0.86 0.96 1.29
27 PeixotoajussieuanaJuss.(Malpighiaceac)
JRL80 2.93 3 3 0.86 0.78 1.29
28 MachaeriumacutifbliionVogei.(Fabaceae) JRL26 2.25 2 2 0.57 0.82 0.86
29 ManihotpalrnataMull.Arg.(Euphorbiaceae)
JRL66 2.24 3 2 0.86 0.52 0.86
30 ErythroxylumlaetevirensO.E.Schulz(Erythroxylaceae)
LWLV1070 1.94 2 2 0.57 0.51 0.86
3 DalbergiacearensisDucke(Fabaceae)
1 JRL43 1.94 2 2 0.57 0.51 0.86
32 Jacarandajasminoides(Thunb.)Sandwith(Bignoniaceae)
JRL79 1.91 2 2 0.57 0.48 0.86
33 CmtongrewioidesBaill.(Euphorbiaceae) JRL86 1.78 2 2 0.57 0.35 0.86
34 ErythroxylumstipulemmPlowman(Erythroxylaceae)
JRL69 1.78 2 2 0.57 0.34 0.86
3356 AEsrpyithdroosxpyelrummacfs.uvbaicccainniuimfoMlairutm.Ma(rAtp.oc(Eyrnyatcheraoex)ylaceae) JRL 17 1.62 1 1 0.29 0.9 0.43
JRL27 1.3 2 1 0.57 0.3 0.43
37 Gymnanthessp2(Euphorbiaceae)
JRL 107 1.25 2 1 0.57 0.25 0.43
38 Arrabidaeasp.(Bignoniaceae)
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386 Lima,J.R.etal.
No Species/ Family Voucher IV N NP ReD ReBa ReF
39 AgonandrabrasiliensisMiers(Opiliaceae) JRL56 1.22 1 1 0.29 0.51 0.43
40 Eugeniacf.piauhiensisO.Berg(Myrtaceae) JRL62 1.22 1 1 0.29 0.51 0.43
41 Secondontiacf.foliosaA.DC.(Apocynaceae) JRL89 1.22 1 1 0.29 0.51 0.43
42 Erythroxylumsp.(Erythroxylaceae) JRL87 1.07 1 1 0.29 0.35 0.43
43 SennacearensisAfran.Fem.(Fabaceae) JRL46 1 1 1 0.29 0.28 0.43
44 PiptadeniamoniliformisBenth.(Fabaceae) FSA1298 0.89 1 1 0.29 0.17 0.43
45 Undackeriaovata(Benth)Gilg(Achariaceae) JRL76 0.84 1 1 0.29 0.13 0.43
46 Sapiumaff.argutum(Midi.Arg.)Huber (Euphorbiaceae) JRL75 0.84 1 1 0.29 0.13 0.43
47 Tocoyenaformosa(Cham.&Schltdl.)K.Schum.(Rubiaceae) JRL59 0.84 1 1 0.29 0.13 0.43
48 Thiloaglaucocarpa(Mart.)Eichler(Combretaceae) LWLV1050 0.84 1 1 0.29 0.13 0.43
49 Ourateasp.(Nyctaginaceae) JRL54 0.84 1 1 0.29 0.13 0.43
50 CordiarufescensA.DC.(Boraginaceae) JRL24 0.84 1 1 0.29 0.13 0.43
Table5-FamiliesandspeciesofherbandsubwoodyplantsintheforestofResevaNaturalSerradasAlmas,Crateús,
Ceará State, with their respective proportions ofground cover and densities. Collectors: FSA - Francisca Soares
Araújo,JRL-JaciraRabeloLima, LWLV-Luis Wilson Lima-Verde.
Family / species Cover% Density plant in 100 m2 Voucher
Araceae
ScaphispathahastifoliaHook. 1 58 FSA 1379
Bromeliaceae
Bromeliaplumieri(E.Morren)L.B. Sm. 1 0 LWLV982
BromeliaauriculataL.B.Sm. 8 79 LWLV 1222
Cyperaceae
Cyperusagregatus(Willd.)Endl. 1 8 JRL 106
Malvaceae
Pavonia sp. 3 168 JRL90
Poaceae
Lasiaciscf.sorghoidea(Desv.exHam.)Hitchc.&Chase 1 33 FSA 1378
Streptostachysasperifolia(Kuntw.)Desv. 18 580 FSA 1307
coupled with differences in methodology. One of has usually been ignored in vegetation studies in
these studies, including WCSP, wasconducted in the region. In fact, its contribution to the basal
the sameplaceasourstudy andalso inCurimataú area is small and it has little value in surveys
(PB), and Betânia (PE) (Araújo et al. 2005a). directed to estimate fuel wood or timber
However, the sampling method was the point availability.Floristicsurveysusuallyconsiderthat
quartermethod,contrary totheplotmethodinour the plants in this component are merely young
study, making it difficult to compare results. individuaisofspecieswhicharealsopresentinthe
Differences shown by the two methods for Serra larger component (WCLP). This occurs in
daAlmas(Tab.7)illustratethisdificulty. forestselsewhere(Whittaker1975;Quigley&Platt
The openness of the canopy in seasonal 2003). However, one family (Asteraceae) was
montaneforestsallowsnotonlytheestablishment exclusive and another one (Verbenaceae) had
but also the maintenance ofthe WCSP (Quigley most of its representatives in this component,
&
Platt2003),evenoutsidegaps(Whittaker 1975; including some exclusive species (Lantana cf.
Gentry & Dodson 1987). The WCSPcomponent brasiliensisandLantanasp.).Therefore,inclusion
Rodriguésia62(2):379-389.201
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1
Physiognomyandstructure ofa seasonaldeciduousforestin Ceará, Brazil 387
Table6-Altitude(Alt),meanannualrainfall(MAR),sampledarea;totalplantdcnsity(TPD),communitybasalarea
(CBA),maximumplantheight(MxH),proportionofplantsabove8mheight(>8),andmean(MD)andmaximumstem
diameters(MxD)offorestandnonforestformationsinsemiaridNortheastBrazil(onlysurveyswithinclusioncriterion
ofPSL>3cm).
Location Alt MAR Area TPD CBA MxH H>8 MD MxD References
(m) (mm) (ha) (pl/ha) (mVha) (m) (%) (cm) (cm)
Forestformations
SerradasAlmas,CE 650 1044 1 5683 47.0 18 11 8.4 65 Thisstudy
Triunfo1,PE 1100 1260 02 6515 56.7 14 8 8.1 102 Ferrazetal. 2003
Triunfo2,PE 900 1066 0.1 3060 46.7 15 12 10.5 60 Ferrazetal. 2003
Pesqueira, PE 1082 681 0.3 4910 67.2 16 21 9.8 72 Correia1996
Jataúba, PE 1020-1120 764 02 4406 49.6 15 12 8.7 80 Moura 1997
SerradeBodopitá,PB - 500 02 3165 3128 - - - - Oliveiraetal.2009
SerradeBodocongó,PB - 500 02 3010 33.19 - - - - Oliveiraetal2009
SerradoMonte,PB - 500 02 4530 33.19 - - - - Oliveiraetal.2009
SerradoCanoió,PB - 500 02 4145 23.25 _ _ - Oliveiraetal.2009
Nonforestformations
Padre Marcos, PI 420 637 0.45 4618 24.2 9 <1 8.1 48 Oliveiraetal. 1997
BaixaFria,CE 750-760 838 0.25 5952 14.2 8 10 5.0 29 Araújoetal.1998
Carrasco, CE 750-760 838 0.25 5722 26.8 13 9 6.5 40 Araújoetal. 1998
CEasptirvoanrdao,,CPIE 75600-7060 688380 0.125 65589267 3119..95 1“1 <_1 5..4 2-7 ALreamúojso&etaRlod1a9i928002
Table7-Resultsobrainedwiththepio,andtheqn.rtcrpoin.nrcthodsfo,lhewoodcomponentofsmallperime,cm
(>3cmand<8.9cm)inthesameareaoftheseasonaldeciduous forestofScmrdasAlmas,CearA
Quarter point
Plot
6
22
Numberoffamilies
8
Numberofspecies 50
1.403
ShannonIndex(nats/plant) 3.26
3423
17500
Absolute density (plant/ha)
0.002
Basalarea(nr/ha) 2.8 0.53
Meanstemdiameter(cm) 1.4
of this component is recommended in future scuopmpproensseinotn,ofittpherohbearbbl/ysucbownolordiybuctoemdpotnoentth,e
tqsruteiuetdseiehaDsinegdnihns,sitththhryeruebirensetgti(ihmoiWsneC.scLhoPim)gp,hoenrbeuttnhtaintostfchaottnhteorffitobrhueetsiltoanrwgatesor ttDhheinrssoiuhtgeyhr(bl9/igsphluta,bnwtwosao/tdme2yr) aacnnoddmpsnoouinltrecinoctvnterwcaeogrmeep(elt3oi0wt%,i)onai.sn
Tbthaheseablashaairlgeaahrewdaaeosnfsrtiahttehylearrogsfemrattlhrlee,esaWlniCdghSsthlPryubacbcooomvmpepoo5nne%enntot.f uWmsieutaahsliulnyrterhmeepeBonrrtatszeidloiffaonrtNhooitsrhtechreofamosprteosrntesegin(otRni,cimhnaarcnduyut1ri9en9cg6ec)n.it
epennteatnrgaltee.dTtohgeetvhegeertwaittiohnt,hereinndfleureinncgeiotfdtihfefiWcuCltLtPo aprleaanstsf/omu2n(dBamrubcohsahiegthael.r2dc0n0s5i;tiRees,isfretomal.1620t0o6)1.587
Rodrlguésià62(2):379-3S9.201
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13 14 15 16 17 lí
1
388
Lima,J.R. etal.
TheIargersize(heightanddiameter)ofplants Acknowledgments
inSerradasAlmasthaninlhenonforestformations
may result from greater water availability. Non- The authors thank the Coordenação de
forest sites tend to have less annual rainfall than Aperfeiçoamento de Pessoal de Nível Superior
Serra das Almas and the topographic position (CAPES), for the scholarship, and the Conselho
favors the forest in Serra das Almas. Most ofthe Nacional de Pesquisa (CNPq), for financing the
carrasco sites are on top of the flat plateau and Wpreoject(EditalUniversal,processo476285/2003-8).
their deep, very sandy soil retains little moisture also thank the Associação Caatinga for
afterrainevents.Someoftheinfiltratedwaterseeps infrastruetureinthefieldwork.
totheslopepositionwherethe forestislocated. In
References
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avoid its influence. Many other fountains estruturadeumremanescentedeflorestaestacionai
punctuatethehundreds-of-kilometerslongeastem semidecíduadeterrasbaixasnoNordestedoBrasil.
slopeoftheIbiapabaplateauandafewofthemare RevistaBrasileiradeBotânica27:463-474.
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highersimilarity in structure ofSerraTalhada and da Caatinga. In: Araújo, F.S.; Rodai. M.J.N. &
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hundredsofkilometersaway,althoughotherfactors biodiversidade do bioma caatinga: suporte a
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Talhada forests may result from selective cutting. ccaoantsienrgvaa:çãsou.poMritneisatéersitoradtoégiMaesioregAimobniaeinsted.e
Localinformationreferstothepreviouspresenceof
Brasília.445p.
smmuaclhlpIlaarngtesrattretehsei1n1t0h0emregsiioten.mTahyerehfilgehctdreensgirtoywtohf Barbo&sa,PeMs.sRo.aV,.;ML.iCm.aR,.R.2B.0;05A.graV,egMe.Ft.a;çãCuonhea,flJo.rPa.
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Oxford. Pp. 106-128.
to the relative openness ofthe also tall but Iarger
asbtuenmdmaencdetorfeessmailnletrhterefeosraesntdssharnudbsthien thhiegnhoenr CorresDieiars,rsaeMnr.atSad.çeão1u9md9e6b.MreeEjssottrdraeudtoau.lrtaiUtnduidaveeveremsgiedPtaeadsçqeãuoFeeiddreaar—amlaPtdEa.e
forestformations.
Pernambuco,Recife.89p.
Rodriguésia62(2):379-3B9.201
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cm L2 13 14 15 16 17