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Evidências do efeito Moran na sincronia populacional: uma demonstração em microcosmo experimental.

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ECOLOGY, BEHAVIOR AND BIONOMICS

Evidências do Efeito Moran na Sincronia Populacional: Uma Demonstração

em Microcosmo Experimental

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F.L.V.B. R

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1PPG em Ecologia & Evolução, ICB, Univ. Federal de Goiás, C. postal 131, 74.001-970, Goiânia, GO 2Depto. Biologia, Univ. Federal de Goiás, Campus Jataí, BR 364, km 192, zona rural, 75.801-615, Jataí, GO

paleo_ribeiro@yahoo.com.br

Neotropical Entomology 36(5):662-669 (2007)

Evidences of Moran Effect in the Population Synchrony: A Demonstration in Experimental Microcosm

ABSTRACT - Three main causes to population synchrony are proposed: exogenous factors, dispersal and

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.(<:25'6'HPRJUDSKLFVWRFKDVWLFLW\HQYLURQPHQWDOVWRFKDVWLFLW\HQYLURQPHQWDOYDULDELOLW\

population dynamic

RESUMO - São propostas três causas principais para sincronia de populações: fatores exógenos, dispersão e

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(2)

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em diferentes escalas espaciais – continental, regional e local (Benton et al./LHEKROGet al.)RQWDLQH

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em metapopulações aumenta com a sincronia (Heino et al.

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fragmentadas no espaço (Earn et al.3RUH[HPSORR risco de extinção pode aumentar se a sincronia ocorrer sem

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Atualmente, são propostas três causas principais para sincronia de populações: fatores exógenos, dispersão e

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regionalmente e que compartilham a mesma estrutura de

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o El-Niño (Post &)RUFKKDPPHUHLQFOXLUDOWRVQtYHLV

GHÀXWXDo}HVFRUUHODFLRQDGDVQRHVSDoR%HQWRQet al01, Koenig 2002, )RQWDLQH & *RQ]DOH]

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(Heino 1998, Lande et al. %M¡UQVWDG 1HVWH

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maior que a esperada caso promovam ciclos uniformes (Ranta

et al%ODVLXVet al%M¡UQVWDGH

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dependentes da densidade (Grenfell et al.D

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Entretanto, esses fatores provavelmente agem de forma

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sendo difícil distinguir seus efeitos separadamente em

SRSXODo}HVVLQFURQL]DGDVQDWXUDOPHQWH%HQWRQet al. Para tanto, alguns experimentos em microcosmos (Benton et al)RQWDLQH *RQ]DOH]WrPVLGRUHDOL]DGRVFRP

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Material e Métodos

Biologia do organismo estudado.$VHVSpFLHVGRJrQHUR Sitophilus são popularmente chamadas de caruncho ou gorgulho dos cereais e estão entre as mais difundidas e

GHVWUXWLYDV SUDJDV SULPiULDV GH FHUHDLV DUPD]HQDGRV QR PXQGR (PERUD D IDPtOLD &XUFXOLRQLGDH DJUXSH PXLWDV SUDJDVDJUtFRODVGHVWUXWLYDVDSHQDVDVHVSpFLHVGHSitophilus

VmRLPSRUWDQWHVFRPRSUDJDVGHJUmRVDUPD]HQDGRV*DOOR

et al

Os adultos de SitophilusVSSPHGHPGHPPD mm de comprimento e, em geral, são de vida longa (quatro

DFLQFRPHVHV$VIrPHDVFKHJDPDGHSRVLWDUGHD RYRVGXUDQWHVHXFLFORGHYLGD2VRYRVVmRLQVHULGRV XPDXPGHQWURGHSHTXHQDVFDYLGDGHVDEHUWDVQRVJUmRV H HFORGHP HP DSUR[LPDGDPHQWH VHLV GLDV 5HHV

As larvas se desenvolvem dentro dos grãos e apresentam

FDQLEDOLVPR &RP LVWR UDUDPHQWH HPHUJH PDLV TXH XP LQGLYtGXRDGXOWRGHXPVLPSOHVJUmRGHWULJRRXGHDUUR] HQTXDQWR GRLV RX WUrV SRGHP HPHUJLU GH XP ~QLFR JUmR GHPLOKR2GHVHQYROYLPHQWRFRPSOHWR±RYRODYDSXSD DGXOWR±pSRVVtYHOHPWHPSHUDWXUDVHQWUHž&Hƒ&H OHYDFHUFDGHGLDVTXDWURVHPDQDVHPFRQGLo}HVyWLPDV aƒ&HGHXPLGDGHUHODWLYD(YDQV2V

indivíduos apresentam aumento de mortalidade em grãos

FRPWHRUGHXPLGDGHDEDL[RGHHJHUDOPHQWHRVRYRV QmRVmRFRORFDGRVHPJUmRVFRPXPLGDGHDEDL[RGH &HQWUHLQDU

Delineamento experimental. Os carunchos foram coletados

HPXPGHSyVLWRGHPLOKRQD)DFXOGDGHGH$JURQRPLDGD 8QLYHUVLGDGH)HGHUDOGH*RLiVGLYLGLGRVHPSRSXODo}HV GHLQGLYtGXRVHFRORFDGRVFDGDJUXSRHPXPUHFLSLHQWH GHEDUURGHFP3FRPJGHPLOKR2VUHFLSLHQWHVIRUDP

ODFUDGRVFRPXPWHFLGRGHPDOKD¿QDGHIRUPDDLPSHGLU

a dispersão dos carunchos entre populações, mas permitir a

SDVVDJHPGHDU

$V SRSXODo}HV XQLGDGHV H[SHULPHQWDLV IRUDP VXEPHWLGDVDGRLVWUDWDPHQWRVGLVWLQWRVFDUDFWHUL]DQGRD YDULDomRDPELHQWDO6HLVSRWHVIRUDPFRORFDGRVQXPDFDL[D GHSDSHOmRIHFKDGDFPGHDOWXUDFPGHODUJXUDH FPGHHVSHVVXUDFRPSHTXHQRVRULItFLRVQDVODWHUDLVVHLV

(3)

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ressaltar que todas as etapas do experimento, tanto de seleção dos indivíduos e recipientes, quanto dos tratamentos aos quais

IRUDPLQVHULGRVRFRUUHUDPGHIRUPDWRWDOPHQWHDOHDWyULD 2FHQVRWRWDOGDDEXQGkQFLDSRSXODFLRQDOIRLUHDOL]DGRD FDGDVHWHGLDVHQWUHHDWUDYpVGHFRQWDJHP GLUHWDGHWRGRVRVLQGLYtGXRVDGXOWRVHPFDGDSRSXODomR

A temperatura e umidade relativa foram registradas em cada tratamento durante todo o experimento, com as caixas

IHFKDGDVXWLOL]DQGRXP7HUPR+LJU{PHWURGLJLWDO0,1,3$

Análise dos dados2VGDGRVRULJLQDLVGHDEXQGkQFLDGDV

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Posteriormente, foram ajustados modelos autoregressivos, separadamente, para os dados temporais de cada população (ver Ranta et al..RHQLJ%XRQDFFRUVLet al. Os resíduos dos modelos autoregressivos de primeira ordem

IRUDPDQDOLVDGRVQRYDPHQWHDWUDYpVGHXPFRUUHORJUDPDFRP RREMHWLYRGHYHUL¿FDUVHDVWHQGrQFLDVGHORQJRSUD]RIRUDP UHPRYLGDVRXVHMDYHUL¿FDUDDXVrQFLDGHDXWRFRUUHODomR HQWUHRVUHVtGXRV

$YDULDomRGRQ~PHURGHLQGLYtGXRVHQWUHDVSRSXODo}HV IRL REWLGD DWUDYpV GR FiOFXOR GR GHVYLR SDGUmR GH VXDV DEXQGkQFLDVGDGRVWUDQVIRUPDGRVSDUDOQSDUDFDGDXQLGDGH WHPSRUDOFROHWDHQXPPHVPRWUDWDPHQWR$PpGLDGHVVHV

desvios padrão em cada tratamento, considerando toda a

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3DUDDYDOLDUDVLQFURQLDSRSXODFLRQDOIRUDPXWLOL]DGRV

os coeficientes de correlação de Pearson e Spearman (Buonaccorsiet alHQWUHWRGDVDVSRSXODo}HVGHQWUR e entre tratamentos, usando os resíduos dos modelos

DXWRUHJUHVVLYRV4XDQWRPDLRUDVLQFURQLDHQWUHDVÀXWXDo}HV

populacionais, maior o coeficiente de correlação (mais

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sincronia entre as populações dentro e entre tratamentos, foi construído um box-plot FRPDPpGLDHRHUURSDGUmR

GRVFRH¿FLHQWHVGHFRUUHODomR1HVVHFDVRDKLSyWHVHDVHU WHVWDGDpVHDVSRSXODo}HVDSUHVHQWDPPDLRUVLQFURQLDHQWUH VLGRTXHHQWUHWUDWDPHQWRV

2 WHVWH GH 0DQWHO 0DQWHO IRL UHDOL]DGR SDUD YHUL¿FDUDFRUUHODomRGDGLVWkQFLDHVSDFLDOHQWUHDVXQLGDGHV H[SHULPHQWDLVPHQVXUDGDDWUDYpVGDGLVWkQFLDHXFOLGLDQD FRPVXDVUHVSHFWLYDVÀXWXDo}HVQDGHQVLGDGHEDVHDGDQD FRUUHODomRGH3HDUVRQFRPRPHGLGDGDKHWHURJHQHLGDGH HVSDFLDOQRWUDWDPHQWRFRPOkPSDGD2WHVWHGH0DQWHO= pGDGRSRU

= ™;ij Yij, n

i, j = 1

onde, Xije YijVmRRVHOHPHQWRVGDVPDWUL]HV;H<DVHUHP

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$V YDULiYHLV DPELHQWDLV WHPSHUDWXUD H XPLGDGH TXDQWL¿FDGDVQRH[SHULPHQWRIRUDPFRUUHODFLRQDGDVHQWUH VLGHQWURHHQWUHWUDWDPHQWRVFRPD¿QDOLGDGHGHVRQGDUD LQÀXrQFLDGRVIDWRUHVDPELHQWDLVSRVVLYHOPHQWHHQYROYLGRV QRVUHVXOWDGRVHQFRQWUDGRV(VVDVYDULiYHLVWDPEpPIRUDP DQDOLVDGDVDWUDYpVGHXPFRUUHORJUDPDSDUDLGHQWL¿FDUR SDGUmRDOHDWyULRRXFtFOLFRGDVRVFLODo}HVQDWHPSHUDWXUD HXPLGDGH

$ VLJQLILFkQFLD HVWDWtVWLFD GH WRGDV DV DQiOLVHV GH FRPSDUDomRHQWUHPpGLDVIRLREWLGDSHORWHVWHWGH6WXGHQW HSHOD$QiOLVHGH9DULkQFLDSDUDFHQiULRVFRPGXDVPpGLDV HPDLVGHGXDVPpGLDVUHVSHFWLYDPHQWH

Resultados

Os dois tratamentos foram mantidos durante o tempo planejado e nenhuma população se extinguiu no decorrer

GR H[SHULPHQWR 1R HQWDQWR DSHVDU GH RV FDUXQFKRV DSUHVHQWDUHPHPJHUDOWHQGrQFLDFUHVFHQWHQRQ~PHURGH

indivíduos, existe clara diferença nas taxas de crescimento

HQWUH DV SRSXODo}HV FRQWLGDV QRV GRLV DPELHQWHV VHP OkPSDGDLQFUHPHQWRPpGLRGHLQGGLDFRPOkPSDGD LQFUHPHQWR PpGLR GH LQGGLD (VSHFL¿FDPHQWH QR DPELHQWHVHPOkPSDGDDVSRSXODo}HVDSUHVHQWDUDPWD[DV GHFUHVFLPHQWRVLJQL¿FDQWHPHQWHPDLRUHVHPDLVFRQVWDQWHV TXHQRDPELHQWHFRPOkPSDGDW 3

$VSRSXODo}HVPDQWLGDVFRPOkPSDGDDSUHVHQWDUDPPDLRU YDULDomRQRQ~PHURGHLQGLYtGXRVDRORQJRGRWHPSRPpGLD GRGHVYLRSDGUmRGDDEXQGkQFLDDFDGDXQLGDGHWHPSRUDO ±WUDWDPHQWRFRPOkPSDGDVPpGLR WUDWDPHQWRVHP OkPSDGDVPpGLR W 3FRPIDVHVGH GHFOtQLRLQWHUFDODGDVFRPFUHVFLPHQWRSRSXODFLRQDO)LJD $SHVDUGHHVVDYDULDomRVHUPDLRUQRWUDWDPHQWRFRPOkPSDGD RFRUUHORJUDPDGRVGDGRVRULJLQDLVGHDEXQGkQFLDLQGLFD

um claro padrão de autocorrelação temporal para todas as

SRSXODo}HVHPDPERVWUDWDPHQWRVQRVSULPHLURVKRUL]RQWHV WHPSRUDLV)LJELQGLFDQGRDWHQGrQFLDGRFUHVFLPHQWR SRSXODFLRQDODRORQJRGRWHPSRMiLGHQWL¿FDGDQD)LJD

2FRUUHORJUDPDUHSUHVHQWDGRQD)LJLQGLFDDXVrQFLD

de autocorrelação temporal nos resíduos após o ajuste de

PRGHORVDXWRUUHJUHVVLYRVWRUQDQGRVHHYLGHQWHDVÀXWXDo}HV

na densidade de cada população com a remoção da tendência

GHFUHVFLPHQWRGHORQJRSUD]R

2VFRH¿FLHQWHVGHFRUUHODomRGH3HDUVRQH6SHDUPDQ

mostraram-se maiores entre as populações de mesmo tratamento do que entre as populações de tratamentos

(4)

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nesse estudo correlacionaram-se de forma positiva entre os

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e negativa entre si no mesmo tratamento (temperatura

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WUDWDPHQWRVHPOkPSDGD“o&W 3 enquanto a umidade apresentou níveis mais elevados na

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apresentaram tendência de crescimento ao longo do tempo,

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GHXPDSRSXODomR

0 5 10 15 20 25

Tempo (semanas) 0

1 2 3 4 5 6 7

Abundância (ln)

sem lâmpada com lâmpada

a

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15

Horizonte -1,0

-0,8 -0,6 -0,4 -0,2 0 0,2 0,4 0,6 0,8 1,0

Autocorrelação

sem lâmpada com lâmpada

(5)

quantificadas no experimento (temperatura e umidade

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tratamento não apresentaram autocorrelação temporal e

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ocorreram de forma aleatória e sincrônica entre tratamentos,

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contínuas normalmente levam as populações a ser menos

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e história evolutiva dos organismos (Trenham et al Byholmet al/LHEKROGet al.

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populacional em carunchos nesse sistema experimental, uma

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0 2 4 6 8 10 12 14 16

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Autocorrelação

sem lâmpada

com lâmpada

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SC SS CC

0 0,1 0,2 0,3 0,4 0,5 0,6 0,7 0,8 0,9

Coeficiente de correlação

Pearson Spearman

média ± erro padrão

(6)

De acordo com o delineamento experimental, as populações

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naturais, enquanto as demais foram expostas a condições

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alterações fisiológicas nos organismos (estimulação ou

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et alSDUDSancassania berlesei0LFK$FDULQD

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via mecanismos amplamente similares nos organismos,

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de persistência de populações em um sistema experimental aumenta com a autocorrelação espaço-temporal entre as

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das seis populações expostas a regimes diferentes de oscilações

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mantendo-as com densidades mais elevadas e, portanto,

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medida que as populações crescerem, a competição por

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a pressão competitiva entre os indivíduos e, posteriormente,

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semelhantes (mesmo regime de variação na temperatura

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01 04 06 10 13 16 18 22 25 Tempo (semanas)

26 28 30 32 34 36 38 40 42 44 46 48 50

Temperatura (ºC)

0 20 40 60 80 100 120

Umidade relativa (%)

a

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 Horizonte

-0,3 -0,2 -0,1 0,0 0,1 0,2 0,3

Autocorrelação

b

Temp. sem lâmpada Temp. com lâmpada

(7)

flutuar de forma sincrônica (competição interespecífica,

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escalas espaciais, nossos resultados permitiram evidenciar a

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Síntese pela concessão do espaço físico para o experimento,

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