Quaternary Research 56, 383389 (2001)doi:10.1006/qres.2001.2264, available online at http://www.idealibrary.com on

T m

e

Palyngion invegetatlowlandridge vcarbon14C yrduring14C yr BpatchesMoracesent dushiftedtween 3duringoped asB.P. ThAtlanticturbedtil ca. 6foraminstudy si

Key Wforest; A

Paly20050S300 kmland anby tropforest twhichhighlanropical Rain Forest and Climate DynaSouthern Brazil, during the

Hermann BehlingCenter for Tropical Marine Ecology, Fahrenheitstra

E-mail: hbehling@zmt.uni-br

andRaquel R. B. NegrelUniversidade Federal do Parana, Departamento de Botanica,

Received October 18, 2000; published onlin

ological analysis of a core from the Atlantic rain forest re-Brazil provides unprecedented insight into late Quaternaryional and climate dynamics within this southern tropical. The 576-cm-long sediment core is from a former beach-alley, located 3 km inland from the Atlantic Ocean. Radio-

dates suggest that sediment deposition began prior to 35,000B.P. Between ca. 37,500 and ca. 27,500 14C yr B.P. andthe last glacial maximum (LGM; ca. 27,500 to ca. 14,500.P.), the coastal rain forest was replaced by grassland andof cold-adapted forest. Tropical trees, such as Alchornea,

ae/Urticaceae, and Arecaceae, were almost completely ab-ring the LGM. Furthermore, their distributions wereat least 750 km further north, suggesting a cooling be-C and 7C and a strengthening of Antarctic cold fronts

full-glacial times. A depauperate tropical rain forest devel-part of a successional sequence after ca. 12,300 14C yr

ere is no evidence that Araucaria trees occurred in thelowland during glacial times. The rain forest was dis-

by marine incursions during the early Holocene period un-100 14C yr B.P., as indicated by the presence of micro-ifera. A closed Atlantic rain forest then developed at thete. C 2001 University of Washington.ords: Brazil; late Quaternary; pollen; tropical Atlantic rainraucaria forest; grassland.

INTRODUCTION

nological studies from Catas Altas (755 m elevation,, in southeastern Brazil 43220W), at a distance of aboutfrom the Atlantic Ocean, indicate that glacial grass-

d small areas of subtropical gallery forests were replacedical semideciduous forest and cerrado (savanna, savannao woodlands) (Behling and Lichte, 1997). Grasslands,occur today in small patches in the southern Braziliands, expanded at least 750 km northward during the

recordevegetatian avera

markedvationsAtlanticfor the wdecreas(LGM;possiblylowland

Paleocal Atlanothingtions (Brain forhas beeies (e.g.exists toforests,the Atlaa few smsome su

last glacinvestigAtlanticyses ofVelha, S

The sbiologicbiosphenorthern

383ics of the Atlantic Lowland,Late Quaternary

e 6, 28359 Bremen, Germanymen.de

leC.P. 19031, 81531-970 Curitiba, Brazil

e September 5, 2001

d glacial period (ca. 18,000 to>48,000 14C yr B.P.). Theon reconstructions suggest a dry and cold climate withge annual temperature reduction of 57C. Did thesechanges in climate, as documented from the lower ele-in the highlands, also affect the tropical rain forest of the

lowland? Estimated sea-surface temperatures (SSTs)estern tropical South Atlantic Ocean show only a minor

e in temperature (2C) for the last glacial maximumCLIMAP, 1981). Such a small change in temperaturehad only minor consequences on the distribution of therain forest.environmental data from the region of modern tropi-ntic rain forest are only of Holocene age, and virtuallyis known about glacial, especially full-glacial, condi-

ehling, 1995a, 1995b). The fragmentation of the Atlanticest into relatively isolated islands during glacial timesn hypothesized in both zoo- and phytogeographic stud-, Muller, 1973; Prance, 1982), but insufficient evidence

test this idea. It is also unknown whether Araucaria

which today occur in the southern highlands, occupiedntic lowland. The modern occurrence of Araucaria onall isolated hills in the southern Atlantic lowland lends

pport to the regional persistence of these trees during theiation (Reitz and Klein, 1966). We present results of anation into past vegetation and climate in the southernrain forest region, using pollen and radiocarbon anal-recently discovered late Pleistocene deposits at Voltaanta Catarina, Brazil.

STUDY AREA

tudy area (26040S, 48380W) lies within the Volta Velhaal reserve, which is part of the larger Mata Atlanticare reserve. Volta Velha is located in the Itapoa district of

Santa Catarina State (Fig. 1). The study area is only a0033-5894/01 $35.00

Copyright C 2001 by the University of Washington.All rights of reproduction in any form reserved.

384 BEHLING AND NEGRELLE

e

few mAtlantain.plainQuat(Marsmalcorinlong

Thwide(Kledone1995incluing ediaceacipacons

Miconea)1-yemon

polleet al

Thwith no frost1986; Nimer, 1do Sul (ca. 15nual rainfall o21:4C. Joinvirecords valueslute minimum

Sampling andThe deepest

corer. The tota

la

s

f

fge

t)t

r

a

14105 Dark brown, decomposed organic material with plantor annual dry season (Atlas de Santa Catarina,986). The meteorological station at Sao Franciscokm to the south of the study area) has a mean an-f 1875 mm and a mean annual temperature oflle station (ca. 30 km southwest of the study area)of 2171 mm and 20.3C, respectively. The abso-temperature is >1C.

METHODS

Pollen Analysisportion of the basin was sampled using a Russianl length of the core, starting from the compact

remains and rootlets; 34105 cm: less plant remainsand fine sand

105157 Black, highly decomposed organic material with a fewplant remains and no rootlets; 145157 cm: morecompact

157300 Blackdark gray organic-rich clay, fine sand, rare plantremains; 179300 cm: greater clay component,compact

300438 Dark gray fine sandy clay, less in organic material, rareplant remains

438478 Blackdark gray organic rich clay, richer in clay andcompact

478576 Black, completely decomposed organic material576580 Sandy subsurfaceFIG. 1. Map of the Volta Velha Reserva and oth

eters above modern sea level and lies 3 km to the west of thetic Ocean and 25 km to the east of the Serra do Mar Moun-The topography is a flat to smoothly undulating coastal(0.5 and 5 m above modern sea level), that was formed by

ernary sedimentary deposits of fluvial and marine originstin et al., 1988; Souza, 1999). Coastal beach ridges andl active and inactive river channels are also present. Theg site is part of a shallow basin, ca. 20 m wide and 100 m

. We refer to this landform as a former beach-ridge valley.e tropical Atlantic rain forest zone forms a 100 to 200-km-strip from Natal to Porto Alegre along the Brazilian coast

in, 1978; Por, 1992). A detailed floristic inventory wason a 1-hectare plot adjacent to the coring site (Negrelle,

; Salimon and Negrelle, unpublished data). The plotdes 248 identified tree, shrub, and herb species (exclud-piphytes) from 50 families. Tapirira guianensis (Anacar-ae), Aparisthmium cordatum (Euphorbiaceae), and Ocoteahylla (Lauraceae) are the dominant trees. The understory

ists primarly of Geonoma spp., Melastomataceae (e.g.,nia chartaceae, Miconia hymenonervia, Ossaea sangui-

, and Rubiaceae (e.g., Psychotria spp.). The results of aar sample of modern pollen showed that tropical taxa com-in the vegetation had high pollen abundances, whereasn from the more infequent plant types was rare (Behling., 1997).e study area experiences a warm, humid, tropical climate

sandyTablein p(ca. 8bulkuseddatedGraathrou

Winterving s1989mulaclavapreseeachtree t

Depth

0r localities mentioned in the text.

base, is 580 cm. Stratigraphy of the core is given in1. Sections of 50 cm length were extruded on site, wrappedstic film and aluminum foil, and stored under coolC), dark conditions until sampling could be done. Eightamples and one wood sample (70 cm core depth) wereor radiocarbon dating. An additional eight samples wereby accelerator mass spectrometry (AMS) at the Van derf Laboratory of the University of Utrecht and one sampleh the conventional method by Beta Analytic (Table 2).took 58 subsamples (0.5 cm3) at 5-cm, 10-cm, and 20-cmals for pollen analysis. All samples were processed follow-andard pollen analytical techniques (Faegri and Iverson,. To determine the pollen concentration and pollen accu-ion rate, one tablet containing exotic spores of Lycopodiumtum was added to each sample. Pollen and spores were wellved and a minimum of 300 pollen grains was counted forsample. The total pollen sum includes herb, shrub, andxa. Pollen identification was based on Behlings reference

TABLE 1Stratigraphy of the Volta Velha Core

(cm) Description

14 Brown, relatively well-preserved organic material withplant remains and rootlets

BRAZILIAN RAIN FOREST 385

TABLE 2AMS and Conventional Radiocarbon Dates from Volta Velha Core

Lab num

UtC-549UtC-549UtC-549UtC-550UtC-550UtC-550UtC-550UtC-550Beta-79

Con

collectmorph

Calcand spCONIvidualtypes ((Fig. 3zonatiassem

Radioc

RadVoltaPleistoges (seyr B.P21,310the co14C yrRadiocHoloction wmost l(ca. 47early tbly) in5645model

Volta V

Poll178 poand 3)trationyr) varber Depth (cm) Age (14C yr B.P.) 13C/12C

7 30 3816 36 29:68 70 (wood) 3343 38 30:89 180 6925 44 25:90 280 7600 60 24:21 380 7980 50 26:12 430 7525 44 28:33 480 12;330 70 26:44 530 25;650 170 24:0

120 564575 21;310 160 25:1ventional radiocarbon date.

ion (containing about 2000 Brazilian species) and pollenological descriptions (Behling, 1993).ulations, cluster analysis, and illustration of the pollenore data were done using TILIA, TILIAGRAPH, and

SS (Grimm, 1987). The pollen diagrams include indi-percentages of the most abundant pollen and spore

Fig. 2), total pollen concentration and accumulation rate), and percentages of microforaminfera (Fig. 3). The

on of the pollen record is based on changes in the pollenblages and the CONISS analysis.

RESULTS

arbon Data and Chronological Control

iocarbon dates provide the chronological control for theVelha pollen record and indicate deposits of latecene and Holocene age. Reconstructed vegetational chan-e below) suggest that the AMS date of 25,650170 14C. at 530 cm depth is probably correct, whereas the160 14C yr B.P. date is in error. Thus, the base of

re is seeiningly closer to the extrapolated age of 37,640B.P. than to the radiocarbon age of 21,310 14C yr B.P.arbon dates from the late glaciation (480 cm) and early

ene (430, 380, 280, and 180 cm) indicate that sedimenta-as not continuous during this period. However, depositionikely was continuous during the early to mid-Holocene0100 cm). The sedimentation rate was high during the

o mid-Holocene, while sedimentation was low or (proba-complete during the late Holocene. Radiocarbon dates at75 cm, 430 cm, and 70 cm have been excluded from the agewhich is a linear interpolation between radiocarbon date.

elha Pollen Diagram

en assemblages in the Volta Velha core are diverse, withllen and spore types including 21 unknown types (Figs. 2. Six pollen zones were defined (Table 3). Pollen concen-s (grains/cm3) and pollen accumulation rates (grains/cm2/y greatly throughout the core, probably reflecting differentFIG

.2.

Polle

npe

rcen

tage

diag

ram

oft

hem

ajorta

xafro

mth

eVo

ltaVe

lha

core

.

386 BEHLING AND NEGRELLEFIG

.3.

Sum

mar

ypo

llen

diag

ram

from

Volta

Velh

a,sh

owin

gra

dioc

arbo

nda

tes,

stra

tigra

phy,

ecolo

gica

lgro

ups,

tota

lpol

len

sum

s,po

llen

conce

ntr

atio

n,po

llen

accu

mu

latio

n,po

llen

zon

es,an

dth

ecl

uste

ranal

ysis

dend

rogr

am.

BRAZILIAN RAIN FOREST 387

TABLE 3Pollenof Po

Zone

VV-IVV-IIVV-IIIVV-IVVV-VVV-VI

deposiconcen

VV-IIIPoll

taxa (6by AstePaepalMoritzmostlyArecacmanniaand MAraucaoccurs

Herbthan inAmongtages, wand MativelyresenteCrotonslightly

Zonpollenand bymostlypart ofzone).centag

Zon46%) aMost acaceae

nia, antenuifocentagincludiByrsonincreasMarine

Zonpollen

(up to 94%). In particular, pollen percentages of Myrsine (maxi-

lt

lua

e

e

bimu

eyo

a

sno

croy

nm

l

e

nB0y

n

etiiZones Depth, Calculated Radiocarbon Ages, and Numberllen Samples in Each Pollen Zone of the Volta Velha Core

Depth (cm) Age range (14C yr B.P.) No. of samples

575537.5 37,64027,650 8537.5487.5 27,65014,328 10487.5477.5 14,32812,220 2477.5170 12,2206720 24

17055 67204330 8550 43303190 (or modern) 6

tional environments. For example, particularly high pollentrations and accumulation rates are found in pollen zonesand VV-V.

en zone VV-I is characterized by a predominance of herb777%). Poaceae pollen (4152%) dominates, followedraceae and Cyperaceae. Lesser types include Eriocaulon/anthus with a few grains each of Eryngium, Xyris, andia dasiantha. Shrub and tree pollen (2129%) originatefrom Myrtaceae and Melastomataceae. Pollen grains ofeae, Myrsine, Alchornea, Moraceae/Urticaceae, Wein-, Ilex, Symplocos tenuifolia-type, Podocarpus, Celtis,

imosa II occur but are rare. A single pollen grain ofria angustifolia was found. Pollen of Mimosa II and IIIonly in this zone. Percentages of fern spores are ca. 20%.pollen percentages (7691%) are higher in zone VV-IIzone VV-I. Pollen of shrubs and trees is low (819%).the herbs, Poaceae pollen (6477%) retains high percen-hereas Asteraceae and Cyperaceae decrease. Myrtaceae

elastomataceae are the only arboreal taxa that remain rel-high. Percentages of all other shrubs and trees are rep-d only by single grains or are missing. Pollen grains ofoccur only in zones VV-I and VV-II. Fern spores areless well represented than in the previous zone.

e VV-III is characterized by a remarkable decrease in herb(from 84 to 29%), especially of Poaceae (from 74 to 23%),an increase in shrub and tree pollen (from 14 to 62%),of the Symplocos tenuifoliatype (maximum 6%, firstthe zone) and Ilex (maximum 15%, second part of the

Other arboreal taxa continue to show relatively low per-es. Fern spores increase, including the tree fern Cyathea.e VV-IV is marked by low percentages of herb (27nd high percentages of shrub and tree pollen (4769%).rboreal taxa, such as Myrtaceae, Melastomataceae, Are-, Myrsine, Alchornea, Moraceae/Urticaceae, Weinman-d Ilex, are well represented. Other taxa, such as Symplocosliatype Matayba and Podocarpus, display lower per-es. Several types appear in this zone for the first time,ng Hedyosmum, Piper, Pera, Rhizophora, Struthanthus,ima, and Tapiria. Fern spores have high percentages ande markedly (maximum 124%) at the end of this zone.microforaminifera are also found in the sediments.

e VV-V is characterized by another decrease of herb(down to 1%) and by an increase of shrub and tree pollen

mum atmum, Sduring

Zoneprimarithe grocaceae

decreas

Late Pl

Theat the tmaximern anddistancficientltion ofnever fo(Droserdicate mdepositclays fiin the cthe beaclose ppossiblto early140 cmof marito C4Brazilia1985; Amateriaboth slo

Late Pl

Polle14C yrca. 14,5nated bEryngiucommo

recordsin Santagrows ocamposto modlimitedmounta

The sthe camthe end of this zone), Arecaceae, Weinmannia, Hedyos-oanea, and Clusia increase. Fern spores are also higherhis zone. Microforaminifera are now rare or missing.VV-VI is marked by a slight increase in herb pollen,y at the beginning of the zone (maximum 12%). Amongp of shrubs and trees (8093%), percentages of Are-nd Weinmannia pollen increase, while Myrsine pollens.

INTERPRETATION AND DISCUSSION

istocene and Holocene Depositional Environments

each-ridge valley was ca. 575 cm deeper than todaye of initial organic deposition. During the last glacial

m, Atlantic sea level was ca. 120 m lower than mod-the ocean was at least 100 km east of the site (modernis ca. 3 km). The beach-ridge valley probably was suf-moist to have allowed the accumulation and conserva-rganic-rich material during the late Pleistocene. A lakermed in the valley, however. The few aquatic pollen taxa, Utricularia, Ludwigia) that were found in the core in-oist but not fully hygric conditions. The organic-richalso lack evidence of fluvial transport. Organic-rich

e sandy clays, and/or the presence of microforaminiferare document the occurrence of marine incursions intoh-ridge valley during the early Holocene, suggestingximity of the Atlantic Ocean. These marine incursionshave disturbed and eroded portions of the late glacial

Holocene record. No microforaminifera were found at(i.e., younger than 6100 14C yr B.P.), indicating the ende incursions at the site. A sea level maximum of C3:5

is reported from several localities along the eastn coast during the mid-Holocene (e.g., Suguio et al.,ngulo and Lessa, 1997). The accumulation of organicduring the late Holocene (Zone VV-VI) was probably

w and discontinuous.

istocene Vegetational and Climate Dynamics

assemblages from the pre-LGM (ca. 37,500ca. 27,500.P; Zone VV-I), and the LGM periods (ca. 27,5000 14C yr B.P.; Zone VV-II) indicate a vegetation domi-Poaceae. Asteraceae and Eriocaulon/Paepalanthus,

m, Xyris, and Moritzia dasiantha also were relatively. The pollen percentages are similar to those in Holocene

of the campos (grassland) vegetation from the highlandCatarina (Behling, 1995a). Moritzia dasiantha currentlynly in the highlands. The Volta Velha data suggest thatvegetation, perhaps with a floristic composition similarrn, once occupied the lowlands. Today the campos iso elevations higher than 1000 m in the south Brazilianns.milarity of the LGM assemblage to modern spectra frompos region in the highland of Santa Catarina (Table 4)

388 BEHLING AND NEGRELLE

TABLE 4Last Glacial Maximum Climate Estimates for Volta Velha Based on Comparison between Modern Climatic

( a

o(

n

suggelowerature30 dalower

ThmodeArecalmomodeexam

Urugat leationathe Pca. 2tempnual

ThmataIlex,in thperceoccupangubecauperioonly

ThablyIlex rpenedvalues, indicating a stepwise change toward a warmer climate.These pollen resstage before the eical southern AtVV-IV).

Holocene Vegeta

The compositconstant with on

cne

e

a

estao

,h

et

th

h

(pu

fo

CONCLUSION

e oftudyfor-into,500withand

treesults suggest the occurrence of a successionalxpansion and developement of the diverse trop-lantic rain forest of the early Holocene (Zone

tional and Climate Dynamics

ion of the Atlantic rain forest was relativelyly minor floristic fluctuations from the early

The palynological results document the predominancgrassland-type vegetation during the late Pleistocene at the sarea in the southern Atlantic lowland, where tropical rainest currently exists. The late Pleistocene is sub dividedthree distinct intervals. The pre-LGM (ca. 37,500 and ca. 2714C yr B.P., zone VV-I) is characterized by grasslanda few stands of cold-adapted forests (e.g., MyrtaceaeMelastomataceae) and with rare occurrences of tropicalConditions of Volta Velha, Santa Catarina

Campos region in Pelotas in RiClimate Santa Catarina (1) Grande do Sul

Average temperature 14 to 16C 17.5CMinimum temperature 4 to 8C 2:3CDays of frost per year 20 to 30 days 5 daysAnnual rainfall 1400 mm 1310 mm

a Modern climate datas from Nimer (1986) and Atlas de Santa Catari

sts that the average temperature was about 5:4 to 7:4Cthan today at Volta Velha. The absolute minimum temper-may have been between 4 to 8C, including ca. 20 toys of frost. Annual precipitation perhaps was ca. 475 mmthan present during the full glacial period.

e pollen results show that tropical trees typical of thern Atlantic rain forest (such as Alchornea, Moraceae, and

aceae) were rare in the lowlands during the pre-LGM andst absent during the LGM. Some tropical trees have theirrn limits some hundreds of kilometers farther south. Forple, the present-day range of Alchornea is to the Brazil anduay border (Reitz et al., 1983). Thus, tropical trees foundst 750 km to the north of their current southern distribu-l limit during the LGM. Using meteorological data fromelotas station (Table 4), average annual temperature was:9C lower during the full glacial period. The minimumerature was ca. 2:3C, including 5 days of frost. The an-rainfall would have been ca. 565 mm lower than today.e relatively frequent occurrence of Myrtaceae and Melasto-ceae and of rarer trees, such as Podocarpus, Symplocos, andsuggests the existence of cold-adapted subtropical forestse lowland during the pre-LGM and the LGM. The highntages of herb pollen indicate that only small areas of forestied the lowland, especially during the LGM. Araucaria

stifolia apparently did not grow in the Atlantic lowlandse a single pollen grain was found for the entire glacial

d. Pollen grains of Mimosa II and Mimosa III occurredduring the glacial periods.e late-glacial record (Zone VV-III) at Volta Velha is prob-incomplete. However, Symplocos tenuifoliatype and theneached maximum pollen percentages. This change hap-somewhat before most of the tropical taxa reached higher

Holocludisuggperceacea

the efloodlevelvegecal ptotaltionsalso

DuWeinfrequbeenWith6100Atlan

DuquenThezone

lectswettearea

of TaThemann

rainSite 1), and Pelotas (Site 2)

Climatic differences2) Volta Velha based on site 1=2

21.4C 5.4 to 7:4C/2:9C1C 5C to 9C/3:3C

0 days 2030 days/5 days1875 mm 475 mm/565 mm

a (1986).

ene (Zone VV-IV) until ca. 6700 14C yr B.P. Ferns, in-g some tree ferns, became frequent in the rain forest,sting a wetter climate than previously. Relatively highntages of herb pollen (Poaceae, Cyperaceae, and Aster-) indicate that there was not complete forest cover duringrly Holocene. Herb-dominated vegetation occurred neard lowlands, a result of possible marine incursions. A fewwith single grains of Rhizophora imply that mangrovetion was poorly developed near the study site. Today, lo-pulations of Rhizophora can produce up to 8090% of thepollen sum (Behling et al., 2001). Drier climatic condi-as recorded in the highland (Behling, 1995a, 1997), mayave influenced the vegetation communities.ring the mid-Holocene (Zone VV-V), Myrsine, Arecaceae,mannia, Hedyosmum, Sloanea, and Clusia became morent in the rain forest. Wetter climatic conditions may havehe primary factor in altering the vegetational composition.the marine regression (beginning of Zone VV-V) at about14C yr B.P., open areas became forested, forming a densetic rain forest.ring the late Holocene (Zone VV-VI) Myrsine was less fre-, while Arecaceae and Weinmannia became more frequent.igher occurrence of Poaceae pollen at the beginning of themay suggest some opening of the forest that perhaps ref-uman activity. The increase of Arecacea perhaps indicates

r climatic conditions. Modern pollen rain from the studyBehling et al., 1997) shows a relatively high average valueirira (25%) and low average value of Weinmannia (1.6%).ppermost core sample has 10% Tapirira and 18% Wein-ia pollen, suggesting that pollen from the modern Atlanticrest was not deposited in the uppermost core sediments.

BRAZILIAN RAIN FOREST 389

(e.g., Arecaceae, Alchornea, and Moraceae/Urticaceae). TheLGM (ca. 27,500 and ca. 14,500 14C yr B.P., zone VV-II) hadwidesprcold-adatrees. Texpansistages o

The poccurren

tion typerina. Tro750 kmAtlanticdiversity

CompPelotasgion, wfor the2 to nual prepresent-disagreewere es

(CLIMAPolle

angustifHoweve(e.g., M

MarinHolocenably thepletelyremainetimes. Swith abAlchornsite.

We graour resear

a postdocsupervisiofunded byPalynologthe kind uDeForestthank Rodfirst authoscholarshi

Angulo, RreviewMarine

Atlas de Santa Catarina. (1986). Gabinete de Planejamento e CoordenacaoGeral, Rio de Janeiro. [In Portugese]

ta6H

yH

S,H

a

iHe

.Ht

.Hia

e

.C

EaitM

d

i

.

p,m

t,

.

aDm

gb

, a[I,l

Lo

p3

iciUead grassland with probably only a few small stands ofpted forests and an almost complete absence of tropical

he late-glacial period (zone VV-III) was marked by anon of tropical shrubs and trees, including successionalf Symplocos and Ilex.alynological data from the LGM suggest the widespreadce in the lowland of grassland (campos). This vegeta-is found today in some highland regions of Santa Cata-pical trees, such as Alchornea, were dislocated at leastto the north of their modern distributions. The tropicalrain forest was markedly reduced in both extent and, especially during the full glacial period.aring the modern climate of the campos region and

(southern limit of Alchornea) with that in the study re-e derive a 3 to 7C lower average annual temperatureLGM period. The minimum temperature was between

8C, including 5 to 30 days of frost per year. The an-cipitation may have been about 500 mm lower than theday rainfall. A LGM cooling of at least 3 to 7C is inment with the 2C lower sea-surface temperatures thattimated for the western tropical South Atlantic OceanP, 1981).

n analytical data indicate that there were no Araucariaolia trees in the study region during the late Pleistocene.r, other species that today are restricted to the highlandsoritzia dasiantha) did expand into the lowlands.e incursions affected the study site from the earlye until 6100 14C yr B.P. This marine influence is prob-primary reason the Atlantic rain forest was not com-

closed. Marine incursions notwithstanding, mangrovesd poorly developed near the study site until modernince the mid-Holocene, a dense Atlantic rain forestundant Myrtaceae, Arecaceae, Moraceae/Urticaceae,ea, Myrsine, and Weinmannia has grown at the study

ACKNOWLEDGMENTS

tefully acknowledge the Machado family for allowing us to conductch in their reserve. Field work (1996) was done while H. Behling wastoral fellow at the Smithsonian Tropical Research Institute under then of Dr. Paul A. Colinvaux. Field work and one radiocarbon date wasthe National Science Foundation (BSR-9007019). The Institute of

y and Quaternary Sciences (University of Gottingen) is thanked forse of their pollen laboratory. We thank Dr. Martin Lichte and Dr. HughSafford and Patricia M. Anderson for reading the English text. Weolfo J. Angulo and Dov Por for providing constructive reviews. Ther thanks the Deutsche Forschungsgemeinschaft (DFG) for a researchp and for the financial support of radiocarbon dating.

REFERENCES

. J., and Lessa, G. C. (1997). The Brazilian sea-level curves: A criticalwith emphasis on the curves from the Paranagua and Cananeia regions.Geology 140, 141166.

Behling,VegetaAraukicae 20

Behling,historyHistor

Behling,Pires,ogy 14

Behling,AraucS Braz

Behling,groveBrazil

Behling,rain daBrazil

Behling,at glac

CLIMAPsurfac

Faegri, KWiley,

Grimm,constrCompu

Klein, R.Ilustra

Martin, LgeologSerie G

Muller, PNeotro

Negrellee dinaMun. IParana

Nimer, EGeogr

Por, F.Acade

Prance, GBioloColum

Reitz, R.Itaja.

Reitz, R.Sul. Se

Suguio, KJ. M.nivel dsus im15, 27

Souza, MdinamSubsdTerra,H. (1993). Untersuchungen zur spatpleistozanen und holozanenions- und Klimageschichte der tropischen Kustenwalder und derrienwalder in Santa Catarina (Sudbrasilien). Dissertationes Botan-. J. Cramer, Berlin. [In German]. (1995a). Investigations into the Late Pleistocene and Holocene

of vegetation and climate in Santa Catarina (S Brazil). Vegetationand Archaeobotany 4, 127152.. (1995b). A high resolution Holocene pollen record from Lago do

E Brazil: Vegetation, climate and fire history. Journal of Paleolimnol-253268.. (1997). Late Quaternary vegetation, climate and fire history in the

ria forest and campos region from Serra Campos Gerais (Parana),l. Review of Palaeobotany and Palynology 97, 109121.., Cohen, M. C. L., and Lara, R. J. (2001). Studies on Holocene man-

cosystem dynamics of the Braganca Peninsula in northeastern Para,Palaeogeography, Palaeoclimatology, Palaeoecology 167, 225242.

., Negrelle, R. R. B., and Colinvaux, P. A. (1997). Modern pollena from the tropical Atlantic rain forest, Reserva Volta Velha, SouthReview of Palaeobotany and Palynology 97, 287299.., and Lichte, M. (1997). Evidence of dry and cold climatic conditionsl times in tropical SE Brazil. Quaternary Research 48, 348358.Project Members. (1981). Seasonal reconstructions of the Earthsat the last glacial maximum. GSA Map and Chart Ser., MC-36, 118., and Iversen, J. (1989). Textbook of Pollen Analysis (4th ed.).hichester.

. C. (1987). CONISS: A Fortran 77 program for stratigraphicallyned cluster analysis by the method of the incremental sum of squares.er and Geosciences 13, 1335.

. (1978). Mapa fitogeografico do estado de Santa Catarina. Floraa Catarinense. Herbario Barbosa Rodrigues, Itaja. [In Portugese]., Suguio, K., Flexor, J. M., and Azevedo, A. E. G. (1988). Mapaco do Quaternario costeiro dos Estados do Parana e Santa Catarina.eologia 28, Secao Geologia Basica 18, 140. [In Portugese]

(1973). The Dispersal Centres of Terrestrial Vertebrates in theical Realm. Biogeographica, Vol. II. W. Junk B.V., The Hague.R. R. B. (1995). Composicao florstica, estrutura fitosociologicaica de regeneracao da floresta Atlantica na reserva Volta Velha,

apoa, SC. Unpublished Ph.D. dissertation, Universidade Federal doCuritiba. [In Portugese](1989). Climatologia do Brasil. Fundacao Instituto Brasileiro de

fia e Estatstica, Rio de Janeiro. [In Portugese]. (1992). Sooretama the Atlantic Rain Forest of Brazil. SPBic Publishing, The Hague.. T. (1982). Forest refuges evidence from woody angiosperms. Inical Diversification in the Tropics (G. T. Prance, Ed.), pp. 137156.ia University Press, New York.nd Klein, R. M. (1966). Araucariaceas. Flora Ilustrada Catarinense,n Portugese]Klein, R. M., and Reis, A. (1983). Projeto madeira do Rio Grande dolowia 34-35, 1525. [In Portugese]., Martin, L., Bittencourt, A. C., Bittencourt, A. C. S. P., Dominguez,., Flexor, J. M., and De Azevedo, A. E. G. (1985). Fluctuacoes domar durante o Quaternario superior ao longo do litoral Brasileiro e

licacoes na sedimentacao costeiro. Revista Brasileira de Geociencias286. [In Portugese]. C. (1999). Mapeamento da plancie costeira e morfologia ea das praias do municpio de Itapoa, Estado de Santa Catarina:os a` ocupacao. Unpublished masters thesis, Setor de Ciencias daniversidade Federal do Parana, Curitiba. [In Portugese]

INTRODUCTIONSTUDY AREAFIG. 1.

METHODSTABLE 1TABLE 2FIG. 2.FIG. 3.

RESULTSTABLE 3

INTERPRETATION AND DISCUSSIONTABLE 4

CONCLUSIONACKNOWLEDGMENTSREFERENCES