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Quaternary Science Reviews 19 (2000) 981}994 Late Quaternary vegetational and climate dynamics in northeastern Brazil, inferences from marine core GeoB 3104-1 Hermann Behling *, Helge W. Arz, JuK rgen PaK tzold, Gerold Wefer Center for Tropical Marine Ecology, Fahrenheitstra}e 1, 28359 Bremen, Germany Geoscience Department,University of Bremen, Klagenfurter Strasse, 28359 Bremen, Germany Abstract Late Quaternary paleoenvironments from northeastern (NE) Brazil have been studied by pollen analysis of marine sediment. The studied core GeoB 3104-1 (3340 S, 37343 W, 767 m b.s.l.) from the upper continental slope o! NE Brazil is 517 cm long and '42,000 C yr BP old. Chronological control was obtained by 12 radiocarbon (AMS) dates from individuals of the foraminiferal species Globigerinoides sacculifer. Modern pollen analogs were received from 15 river, lake and forest soil surface samples from NE Brazil. Marine pollen dates indicate the predominance of semi-arid caatinga vegetation in NE Brazil during the recorded period between '42,000 and 8500 C yr BP. The increased #uvial input of terrigenous material, with high concentrations of pollen and specially fern spores, into the marine deposits, about 40,000, 33,000 and 24,000 C yr BP and between 15,500 and 11,800 C yr BP, indicate short-term periods of strong rainfall on the NE Brazilian continent. The expansion of mountain, #oodplain and gallery forests characterize the interval between 15,500 and 11,800 C yr BP as the wettest recorded period in NE Brazil, which allowed #oristic exchanges between Atlantic rain forest and Amazonian rain forest, and vice versa. The paleodata from core GeoB 3104-1 con"rm the, in general, dry pre-Last Glacial Maximum (LGM) and LGM conditions and the change to wet Lateglacial environments in tropical South America. The annual movement of the intertropical convergence zone over NE Brazil, the strong in#uence of the Antarctic cold fronts and changes of the high-pressure cell over the southern Atlantic, may explain the very wet Lateglacial period in NE Brazil. The documented NE Brazilian short-term signals correlate with the documented DansgaardOeschger cycles and Heinrich events from the northern Hemisphere and suggest strong teleconnections. 2000 Elsevier Science Ltd. All rights reserved. 1. Introduction In South America, the region of northeastern (NE) Brazil is still a white spot with regard to the paleoenvironmental history based on pollen analysis. NE Brazil is well known for the occurrence of anomalous, strong droughts (Hastenrath, 1990; Nimer, 1989), which have strong social and economic consequences, so that a study of the history of past environmental changes and their causes is important. Pollen in marine deposits can be an important archive to reconstruct past terrestrial vegetational and climate changes, in view of the di$culties in getting long terrestrial records from lakes, especially as strong droughts can destroy pollen in lacustrine deposits. Pollen analysis of marine sediment cores can contribute * Corresponding author. Tel.: #49-0421-220-8321; fax: #49-0421220-8330. E-mail address: [email protected] (H. Behling). signi"cantly to the understanding of past vegetational and climatic changes on the South American continent as is well demonstrated by studies of the Amazon fan deposits (Haberle, 1997; Hoorn, 1997). Data on Late Quaternary paleoenvironmental changes from NE Brazil are also important for a better understanding of the natural history of the neotropics (Absy et al., 1991; Behling and Lichte, 1997; Behling and Hooghiemstra, 1998,2000; Colinvaux et al., 1996; Ledru et al., 1998; Martin et al., 1997; Van der Hammen and Absy, 1994). Further, from the biogeographic point of view NE Brazil is an interesting region, concerning the origin and past #oristic connections between the Amazonian and Atlantic rain forest areas (Andrade-Lima, 1982; Cole, 1960; Prance, 1985). In order to investigate vegetational and climatic changes in NE Brazil, the pollen and spore content of marine core GeoB 3104-1 was studied. Modern pollen spectra was collected from terrestrial surface samples, from river and lake sediments and forest soils, in order to create a calibration data set. 0277-3791/00/$ - see front matter 2000 Elsevier Science Ltd. All rights reserved. PII: S 0 2 7 7 - 3 7 9 1 ( 9 9 ) 0 0 0 4 6 - 3 982 H. Behling et al. / Quaternary Science Reviews 19 (2000) 981}994 2. Study region 2.1. Modern physical setting The location of the studied marine core GeoB 3104-1 (3340 S, 37343 W, 767 b.s.l.) is 90 km east of the city Fortaleza (CearaH State) on the upper continental slope o! NE Brazil (Fig. 1). The nearest distance to the modern coast line is about 70 km. The continental shelf is here relatively narrow and shallow, 30 km wide and 50 } 80 m deep, and has a steep continental slope (Ponte and Asmus, 1978). The most important source for the #uvial input of terrigenous material into the marine deposits, including pollen and spores, is Rio Jaguaribe, which is about 700 km long and #ows into the Atlantic Ocean about 80 km south of the core site. The river catchment includes the Rio Jaguaribe basin and the )200 km wide relatively #at Atlantic coastal plain, the mountainous hinterland of the Chapada do Araripe, and the Serra dos Cariris Novos between 200 and 1000 m elevation. Trans- portation of pollen and spores by wind to the Atlantic is probably low, because of the main winds blow from the Atlantic Ocean on to the land area (Nimer, 1989). The surface hydrography of the core area is dominated by the northward #owing South Atlantic boundary current (North Brazil Current, NBC). Starting at around 103S as a relatively weak current (North Brazilian Undercurrent " NBUC), the NBC is mainly supplied by South Equatorial Current (SEC) waters and its #ow is closely connected to the shelf edge. A further con#uence with the northern SEC branch at 3}53 S leads to an intensi"cation of the NBC (Condie, 1991; Peterson and Stramma, 1991; Stramma, 1991; Da Silveira et al., 1994). The seasonal variability of NBUC and NBC transport is related to the southeastern trade winds, with a larger transport during austral spring and reduced transport in austral fall (Johns et al., 1998). With annual means of 27.33C, sea surface temperature (SST) shows a seasonal amplitude of ca. 23C. The average sea surface salinity (SSS) is around 36%. Marine pollen transport by the NBC, primarily from the eastern Brazilian coast and with north}northwestward orientation, has to be taken into consideration. 2.2. Vegetation and climate Fig. 1. Location of the marine core GeoB 3104-1 and the terrestrial surface samples from rivers (1}8), lakes (9}10), and forest soils (11}14) in NE Brazil including the schematic surface circulation pattern o! NE Brazil (SEC, South Equatorial Current; NBC, North-Brazil Current; BC, Brazil Current). The modern vegetation in NE Brazil is primary caatinga (IBGE, 1993), which is subdivided into several physiognomic vegetation types, from grassland to xerophytic thorn shrub savanna, and shrub woodland (Cole, 1986; Eiten, 1982). Main taxa belong to families of Mimosaceae, Caesalpinaceae, Fabaceae, Euphorbiaceae, Cactaceae and Poaceae (Hueck, 1966; Sampaio, 1995). Patches of deciduous forest, gallery and #ood plain forests along rivers, and a few small islands of humid forests in mountains over 500 m a.s.l. (Andrade-Lima, 1982) characterize the semi-arid caatinga region. Along the coast di!erent forms of coastal vegetation occur, including the Atlantic rain forest, which is restricted to a small strip along the western coast starting from the city of Natal southwards. Characteristic of the caatinga region is the long dry season (6}11 months), with the main annual rainfall of less than 250}750 mm, occurring between November and March. The average annual temperature is 24}263C. The Atlantic rain forest region re#ects a humid climate with '1250 mm annual precipitation and only a short annual dry period (Nimer, 1989). The general atmospheric circulation of NE Brazil is in#uenced continuously by the southeasterly air #ow from the Atlantic Ocean (Nieuwolt, 1977). These southeasterlies bring relatively dry air masses from the highpressure cell over the southern Atlantic. The rainy season is related to the annual movement of the intertropical convergence zone (ITCZ) and occurs only when the ITCZ moves over NE Brazil. Unusually strong droughts H. Behling et al. / Quaternary Science Reviews 19 (2000) 981}994 occur when the ITCZ remains north of the equator. Depending upon the meteorological conditions these e!ect the behavior of cold fronts from the Southern Hemisphere in eastern and southern Brazil (Ratisbona, 1976). Conditions in the South Atlantic Ocean determine the strength of the subtropical high, the SEC/NBC intensity, and sea surface temperatures (Hastenrath, 1990; Rao et al., 1993; Arz et al., 1998). 3. Material and methods 3.1. Fieldwork The 517 cm long marine gravity core GeoB 3104-1 was collected during the RV VICTOR HENSEN cruise JOPS II, Leg 6 Fortaleza } Recife, in March 1995 (PaK tzold et al., 1996; Arz et al., 1998) from the upper continental slope (water depth: 767 m) o! Fortaleza. Terrestrial surface samples for modern pollen data were collected from rivers, lakes and forest soils in NE Brazil by Dr. Matthias Tintelnot (Senckenberg Institute in Wilhelmshaven, Germany) and by the "rst author (Table 1 and Fig. 1). 3.2. Radiocarbon dates Each C AMS sample consisted of about 700 handpicked individuals of the foraminiferal species Globigerinoides sacculifer (250}400 lm). Carbonate hydrolysis and CO reduction was performed in the Geoscience Department of Bremen, Germany. The AMS measurements were performed at the Leibniz- 983 Table 2 List of AMS-radiocarbon dates from core GeoB 3104-1 Lab. number Depth (cm) C yr B.P. Calibrated Age (cal B.P.) KIA-653 KIA-1857 KIA-1856 GrA-3719 KIA-1855 GrA-3720 KIA-651 GrA-3721 KIA-1853 KIA-1852 KIA-650 GrA-3722 8 20 52 87 97 172 209 274 292 337 384 462 2660$50 5740$60 9660$50 12,580$100 12,960$90 16,120$160 20,540#350/!330 25,050$200 27,820#290/!280 31,690#450/!420 33,400#1840/!149 38,600#900/!800 2880 6450 11,140 14,760 15,230 19,150 24,280 29,340 32,340 36,370 38,100 43,130 Labor in Kiel, Germany (Nadeau et al., 1997) and the Center for Isotope Research, Groningen, The Netherlands (Table 2). 3.3. Stable isotopes Eight specimens of 350}400 lm diameter (measured along the longest axis) of the planktonic foraminiferal species Globigerinoides sacculifer (without "nal chamber) were used for stable isotope measurements. Sample preparation was performed employing an automatic carbonate preparation system attached to a Finnigan MAT 251 mass spectrometer. Analytical internal longtime precision for dO is better than $0.07. Table 1 Location of terrestrial surface samples and the corresponding vegetation units from NE Brazil No. Name Coord. (approx.) State Vegetation 4325N, 37345W 4334N, 37346W (between the sites 1and 4) 5338N, 35323W 5347N, 35315W 10316N, 36332W 14317N, 39300W 15351N, 38355W CearaH CearaH Caatinga Caatinga Caatinga Caatinga Caatinga Cerrado/Caatinga Caatinga/Atlantic rain forest Semideciduous forest/Atlantic rain forest River 1 2 3 7 4 13 12 11 Rio Rio Rio Rio Rio Rio Rio Rio Lake 6 5 Lagoa Grande Lago Bolim 7304N, 41325W 6302N, 35311W PiaumH Rio Grande do Norte Caatinga Restinga Forest soil 8 17 9 15 16 Picos Picos Mirim Campina Grande I Campina Grande I 7304N, 7304N, 5338N, 7313N, 7313N, PiaumH PiaumH Rio Grande do Norte Paraiba Paraiba Caatinga (secondary) Caatinga (secondary) Caatinga (secondary) Deciduous forest Deciduous forest Jaguaribe I Jaguaribe II (unclear) Mirim Potengi Sao Francisco de Contas Jequitinhonha 41325W 41325W 35323W 35348W 35348W Rio Grande do Norte Rio Grande do Norte Sergipe Bahia Bahia 984 H. Behling et al. / Quaternary Science Reviews 19 (2000) 981}994 3.4. Pollen analysis Along the pro"le 52 samples (4 cm and 1 cm thick) were collected at 10 cm intervals for the pollen analysis. All samples, including terrestrial surface samples, were treated with standard acetolysis method, using sodium pyrophosphate and heavy liquid separation (Faegri and Iversen, 1989). Pollen preparation of marine samples included addition of exotic Lycopodium spores to determine pollen concentration (grains/cm) and pollen in#ux (grains/cm/yr). Samples were counted until a minimum of about 300 pollen grains or 100}200 pollen grains for samples with a low pollen concentration. Twelve samples of the core contained insu$cient pollen grains for statistical calculation. Marine micro-foraminifera were also counted on the pollen slides. For identi"cation of pollen grains and spores, morphological descriptions published by Behling (1993), Hooghiemstra (1984), Roubik and Moreno (1991), Salgado-Labouriau (1973) and Behling's own reference collection were used. The total pollen sum, on which the percentage calculation is based, includes herbs, shrubs and trees, but not, fern spores, moss spores and microforaminifera. A few reworked pollen grains and spores were not counted. Identi"ed pollen and spore taxa have been preliminary grouped into herbs, shrubs and trees, exotic pollen taxa, ferns and mosses. The "nal grouping of identi"ed taxa into di!erent ecological groups such as caatinga, Atlantic rain forest, gallery and #ood plain forests, mountain vegetation has not been "nished, because of lacking pollen records and additional surface samples from di!erent vegetation forms in NE Brazil. TILIA, TILIAGRAPH and CONISS software was used to plot the pollen data and to make calculations and cluster analysis (Grimm, 1987). Downcore changes of the most frequent and important pollen and spore taxa are illustrated in the form of a pollen percentage diagram, a summary percentage diagram. The latter record includes also radiocarbon dates, the sum of microforaminifera, pollen concentration, pollen in#ux record and a cluster analysis dendrogram. Fig. 2. Age-depth curve for core GeoB 3104-1. rigenous input (Tintelnot, 1997; Arz et al., 1998, 1999). Downcore variations in the sediment composition re#ect repeated periods of increased terrigenous sediment input which dilutes the marine biogenic signal (carbonate sedimentation). Thus, sedimentation rates are generally high (&14.5 cm/kyr), favoring a high-resolution paleoclimatic study (Fig. 3). 4.2. Oxygen isotope data (Fig. 3) G. sacculifer is a planktonic foraminifera living in the uppermost 80 m of the water column. Besides the e!ect of the glacio-eustatic sea-level change } the so called `ice e!ecta } its dO signal re#ects local variations in surface water temperature and salinity. The global Holocene/LGM amplitude in dO of 1.2 is exceeded in our isotope curve by about 0.7. Oxygen Isotope Stages (OIS) 3 and 2 are characterized by several distinct oscillations with amplitudes as much as 0.6. These isotopic shifts appears to line up with the changes in the sediment composition (Fig. 2). 4.3. Modern terrestrial pollen data (Fig. 1, Table 1, Fig. 4) 4. Results 4.1. Stratigraphy (Fig. 2, Table 2) The chronostratigraphy of core GeoB 3104-1 is based on linearly interpolated C AMS dates (Table 2 and Fig. 2). The AMS dates were corrected by a reservoir age of 400 yr (Bard, 1988). The radiocarbon stratigraphy is in good agreement with the marine oxygen isotope stratigraphy (Fig. 3). During the Last Glacial period and the deglaciation period, sedimentation processes on the upper continental slope o! Brazil were strongly in#uenced by riverine ter- Surface samples from rivers (Nos. 1}8), lakes (Nos. 9}10) and forest soils (Nos. 11}15) in NE Brazil (Fig. 1, Table 1) show the pollen representation of the modern vegetation (Fig. 4). We have to consider the human in#uence on the vegetation when using these modern pollen analogs. Di!erent herb pollen taxa, primarily Poaceae, Cyperaceae, Borreria, Asteraceae are predominant in the surface sediments. Pollen grains of arboreal taxa, such as Melastomataceae/Combretaceae, Arecaceae (Palmae), Moraceae/Urticaceae, Mimosa and Fabaceae are less frequent, except two forest soil samples (Nos. 11 and 12) which show a strong local input of Mimosa. Remarkable is the presence of Hedyosmum pollen in one H. Behling et al. / Quaternary Science Reviews 19 (2000) 981}994 985 Fig. 3. Oxygen isotope curve (G. sacculifer) and sedimentation rates (see also Arz et al., 1998) from GeoB 3104-1. The marine isotope stages are indicated at the top. C AMS dating levels are marked with black arrows. forest sample near Picos in PiaumH State. Hedyosmum is not known from NE Brazil. Fern spores seems to be extremely rare in the caatinga region. It is interesting to note, that the presence of Borreria pollen grains (indicator for open vegetation types) decreases in the river samples from north to south, while fern spores (indicator for wetter conditions) increases along the eastern coast southwards. Spores of tree ferns, e.g. from Cyathea (indicator for permanent moist conditions), are not found in modern river surface samples from northern NE Brazil, but only in the southernmost samples (Nos. 7 and 8), where tree ferns grow in the humid Atlantic coastal mountains. 4.4. Marine fossil pollen data (Figs. 5}7) About 160 di!erent pollen and spore types, including 20 unknown types, were recognized. Fig. 5 show the most frequent pollen and spore taxa. All pollen assemblages are characterized by abundant herb pollen (60}75%), primarily Poaceae, Cyperaceae, Borreria, Asteraceae. Less frequent are Alternanthera, Amaranthaceae/ Chenopodiaceae, Gomphrena/Pfaza-type, and a number of other types occur in low percentages. Shrub and tree pollen percentages are lower (15}25%) and primarily represented by Melastomataceae/Combretaceae, Arecaceae, Moraceae/Urticaceae, Alchornea, Myrtaceae, Malpighiaceae, Hedyosmum, Mimosa and many other taxa which were found in trace amounts (not shown in the pollen diagrams). Ferns are mainly represented by indeterminate monolete and trilete spores, Anemia, Cyathea and Selaginella. Remarkable is the evidence of the Cyathea horrida-type (previously called Hemitelia), which is not known from NE Brazil. Unclear is the origin of exotic pollen grains primarily Pinus, and Quercus and less frequent Abies, Betula, Corylus, Ephedra, Juglans and Juniperus-type. Long-distance transport by marine currents or the contamination of the upper sediments may be the reason for presence of these pollen grains. Portions of herb, shrub and tree pollen percentages do not show marked changes during the time, except the increase of shrub and trees, mainly by Melastomataceae/ Combretaceae, Alchornea and Hedyosmum during the Lateglacial. Stronger variations are found among the fern spores. Monolete psilate spores show high percentages during the Lateglacial, and only at this period di!erent tree fern taxa from Cyathea are abundant. The Selaginella spore curve illustrates marked #uctuations corresponding to the rates of pollen concentration and in#ux. The calculated pollen concentration (deposition of pollen grains per cm) and in#ux (deposition of pollen grains per cm and year) show strong variations from very low to high values in the core sediments (Figs. 6 and 7). The accumulation rate of terrestrial pollen and spores in marine sediments were very low for statistical counts in eight samples from glacial periods (423, 403, 343, 303, 293, 223, 217 and 173 cm depth) and in four samples of the mid and late Holocene age (33, 23, 13 and 3 cm depth). Pollen concentration and in#ux are low during most of the time, including the LGM, and higher during short glacial periods about 40,000, 33,000 and 24,000 C yr BP. The highest rates occur during the Lateglacial between 15,500 and 11,800 C yr BP. Values from the early Holocene are low. The in#ux of fern spores (Fig. 6) follows the pattern of the pollen in#ux. The in#ux Fig. 4. Pollen percentage diagram, showing the most frequent pollen and spore taxa of modern terrestrial surface samples from rivers (1}8), lakes (9}10), and forest soils (11}15), in NE Brazil. 986 H. Behling et al. / Quaternary Science Reviews 19 (2000) 981}994 Fig. 5. Pollen percentage diagram, showing the most frequent pollen and spore taxa from the marine core GeoB 3104-1. The pollen sum, on which the percentage calculations are based, includes herbs, shrubs and trees. H. Behling et al. / Quaternary Science Reviews 19 (2000) 981}994 987 Fig. 6. Pollen in#ux diagram, showing the in#ux values of selected herb, shrub and tree taxa, and the in#ux of sums of groups. 988 H. Behling et al. / Quaternary Science Reviews 19 (2000) 981}994 Fig. 7. Summary percentage diagram, showing the sum of the pollen and spore taxa, micro-foraminifera, pollen sum values, pollen concentration and pollen in#ux record, and the cluster analysis dendrogram. H. Behling et al. / Quaternary Science Reviews 19 (2000) 981}994 989 990 H. Behling et al. / Quaternary Science Reviews 19 (2000) 981}994 variation of marine micro-foraminifera (Fig. 6) is not so marked than the in#ux of pollen and spores. 5. Interpretation and discussion 5.1. The marine signal According to the detailed C AMS stratigraphy and oxygen isotope record, core GeoB 3104-1 represents undisturbed and continuously sedimented Last Glacial to Holocene marine deposits. The sediment generally consists of nannofossil foraminiferal ooze (carbonate content of 40 up to 70 wt%) with episodically increasing amounts of terrigenous clays of #uvial origin. Terrigenous input (XRF Ti}Ca ratio) coincides with the short glacial periods of high pollen and spore concentrations about 40,000, 33,000 and 24,000 C yr BP and between 15,500 and 11,800 C yr BP as shown in Fig. 8. During these intervals the oxygen isotopes are most negative, which can be interpreted in terms of increased sea-surface temperatures. Arz et al. (1998) additionally suggested for these periods an increased mixing of the upper water column, accompanied by intensi"ed SE trade winds and accordingly an enhanced north}northwestward transport of the NBC. Although these hydrographic changes are most simply explained by variations in the tropical/subtropical wind "eld, the timing of the signals (based on the calendar year calibrated chronology, Arz et al., 1998) imply strong teleconnections with the short-term climate variations (Daansgaard-Oeschger and Bond cycles) as reported from the North Atlantic high latitudes (Daansgaard et al., 1993; Bond and Lotti, 1995). 5.2. Modern and past pollen sources Pollen grains and fern spores originate from di!erent vegetation types on the NE Brazilian continent, such as from coastal vegetation (Poaceae, Cyperaceae, Borreria), caatinga (Poaceae, Borreria, Mimosaceae), Atlantic rain forest } gallery forest and #ood plain forests (Alchornea, Arecaceae), and humid mountain vegetation (Cyathea) or in general moist forest (ferns and Hedyosmum). The major source of the pollen grains and spores deposited at the site GeoB 3104-1 was apparently the terrigenous #uvial input, most likely from the nearest major river, Rio Juagaribe. This is also indicated by mineralogical studies of core sediments (Tintelnot, 1997). Atmospheric pollen and spore transport was probably low, because the main winds blow from the Atlantic on to the land area. Changes in the rainfall regime on the NE Brazilian continent, caused changes in discharge and in the transport of sediment loads by the river into the Ocean. Modi"cations in the marine currents such as the NBC, probably in#uenced on a smaller scale the #uvial deposits in the marine sediments. In this context it has to be considered, that the continental shelf was exposed during glacial low Atlantic sea-level stands, conditions that favored #uvial transport to the upper continental slope. The high sea-level stand during the Holocene, when marine currents also in#uenced the coastal shelf, apparently diminished terrigenous sediment transport to the slope. This is probably the reason for the very low pollen and spore content of the mid and late Holocene deposits. Pollen and spore concentration and in#ux from the #uvial deposits in the marine sediments re#ect also alternations in the deposition rate of continental sediments. This is in general related to modi"cations in the river discharge by changes in precipitation on the continent. Oscillations of Atlantic sea-level stands, and marine currents may also have in#uenced deposition rates of pollen and spores. However, low deposition rates of pollen and spores suggest low precipitation rates on the NE Brazilian continent during glacial times, including the LGM, while high deposition rates suggest short periods with higher rainfall rates about 40,000, 33,000 and 24,000 C yr BP. The highest precipitation rate is found during the Lateglacial between 15,500 and 11,800 C yr BP. 5.3. Vegetation and climate changes Pollen data from river surface samples, including two from the Rio Jaguaribe itself (Nos. 1 and 2 in Fig. 4), re#ect the modern pollen analogue, which was not possible to obtain from the upper Holocene marine deposits. Those data indicate that herb pollen dominate in the modern human in#uenced open caatinga vegetation. Relatively high percentages of pollen grains of herbs and lower percentages of shrubs and trees from the fossil marine deposits clearly suggest presence of open vegetation types such as caatinga and coastal savanna (abundant Cyperaceae) in NE Brazil during the glacial and early Holocene periods. Abundant taxa such as Borreria, Alternanthera, Amaranthaceae/Chenopodiaceae, and Gomphrena/Pfaza are light tolerant and have strong preferences to vegetation forms with an open canopy (Gentry, 1993). No marked changes between the proportions of herbs, shrubs and trees probably indicate similar semi-arid conditions in NE Brazil during the glacial period than present day. Remarkable are the short periods of higher pollen concentration and in#ux, which show abundant fern spores deposits. There is an interesting gradient in the river surface samples from north to south along the NE Brazilian coast, re#ected in a decrease of Borreria pollen and an increase of fern spores. This gradient re#ects the change from dry vegetation types with an open canopy to closed humid vegetation types. This observation seems to be important for the interpretation of the marine pollen record and suggest periods of moister conditions in NE Brazil. H. Behling et al. / Quaternary Science Reviews 19 (2000) 981}994 991 Fig. 8. Summary of palynological and paleoceanographic data, showing the total pollen in#ux, the fern spore in#ux, the XRF Ti-Ca ratio(scaled up fourfold), and the stable oxygen isotope curve (G. sacculifer) from sediment core GeoB 3104-1. Arrows denote more humid periods in NE Brazil. Selaginella grows on moist soil and may be used as a good disturbance indicator for erosion during strong rain fall. The high presence of Selaginella spores, together with the high deposition rate of pollen, suggest strong rainfall at about 40,000, 33,000 and 24,000 C yr BP under in general dry climatic conditions. These wetter periods were apparently too short for an expansion of humid #ood plain or mountain forests. Only from 15,500 to 11,800 C yr BP is a longer wetter period found which allowed an expansion of humid forests as indicated by the expansion of rain forest (e.g. Alchornea) and humid mountain forests (e.g. Hedyosmum, Cyathea). The annual dry season must have been very short, perhaps less than three months. There was probably a marked expansion of small areas of gallery and #ood plain forests along rivers and humid mountain forests in moist valleys. Hedyosmum, in general an indicator for a cool and humid climate (Reitz, 1965), was apparently frequent in mountain forests and gallery forests along the rivers during the cold Lateglacial period. Tree ferns belonging to Cyathea, which are con"ned to very humid habitats without marked dry season (Golte, 1976), were abundant only during this period and supports the interpretation of a cold and permanent wet climate at that time. Cooling of ca. 53C during the LGM is reported from NE Brazil on the basis of studies on noble gases dissolved in groundwater (Stute et al., 1995). During the early Holocene, Hedyosmum disappears and Cyathea was rare, indicating a return to drier climatic conditions in NE Brazil. Mangrove pollen, especially Rhizophora, were only found in the early Holocene, suggesting that mangrove vegetation was probably rare or absent in the studied coastal region during glacial times. 992 H. Behling et al. / Quaternary Science Reviews 19 (2000) 981}994 5.4. Biogeographic implications During the recorded glacial and early Holocene period, NE Brazil mostly experienced semi-arid climatic conditions as similar to today. However, distinct and short-term wetter periods have been evidenced about 40,000, 33,000 and 24,000 C yr BP and during the longer period of 15,500}11,800 C yr BP. This Lateglacial interval was the wettest period recorded, which allowed marked expansion of moist gallery and mountain forests. During this period #oral migration in both directions between Amazon rain forest and Atlantic rain forest may have taken place. It is possible that during the Lateglacial period a number of present-day humid forest refugia in the mountains (Andrade-Lima, 1982) were connected to form a larger area during cooler and wetter climatic conditions. 5.5. NE Brazil in the context to paleoenvironmental records from adjacent regions Evaluating the new results from NE Brazil, one has to consider that the climate of adjacent regions are in#uenced by di!erent climate regimes. The inferred climatic sequence from core GeoB 3104-1 follows the general trend of dry pre-LGM and LGM conditions, as reported from terrestrial sites in southeastern Brazil such as Catas Altas (Behling and Lichte, 1997) and Morro de Itapeva (Behling, 1997). Periods with relatively wet conditions are inferred from pollen records of Carajas (ca. 40,000}22,800 C yr BP) in southeastern Amazonia (Absy et al., 1991), Aguas Emendadas (26,000}21,500 C yr BP) and CrommH nia (32,000}ca. 20,000 C yr BP) in central Brazil (Barberi, 1994; Ferraz-Vincentini and Salgado-Labouriau, 1996), and Salitre and Serra Negra (40,000}27,000 C yr BP) in southeastern Brazil (Ledru et al., 1996; De Oliveira, 1992). These periods are markedly longer than the wet short-term periods from the record GeoB 3104-1. Continuously wet climatic conditions are reported from central Amazonia (Colinvaux et al., 1996). The wet Lateglacial period between 15,500 and 11,800 C yr BP from NE Brazil is earlier than in the Carajas mountains, which are characterized by forest expansion since around 12,000 until ca. 8,000 C yr BP. The records from central Brazil (Agua Emendadas, and CrommH nia) suggest that dry conditions prevailed (Salgado-Labouriau et al., 1998), but here the dry climate during the early Holocene may have caused the erosion or oxidation of deposits of Lateglacial age. Between 16,000 and 11,000 C yr BP, the Salitre record shows a succession of di!erent forest types, which re#ects increasing moisture. A change from dry LGM conditions to wetter Lateglacial conditions is also reported from Morro de Itapeva in southeastern Brazil. Of interest is the record from Lago do Pires in the Atlantic lowlands of southeastern Brazil (17357 S, 42313 W, 390 a.s.l.) which documents a wetter interval between 8800 to 7500 C yr BP during the markedly dry early Holocene period, (Behling, 1995). 5.6. Possible factors for paleoenvironmental changes in NE Brazil There is a general change from a dry LGM to wetter climatic conditions in tropical South America during the Lateglacial, both north and south of the equator (Behling and Hooghiemstra, 2000). This suggests that modi"cations of the annual movements of the ITCZ between both hemispheres plays an important role in determining past climatic changes. The very wet climatic regime in NE Brazil during the Lateglacial, can be explained by permanent annual movement of the ITCZ over NE Brazil. In their comparative study based on pollen and lake level records in tropical and subtropical South America Martin et al. (1997) conjectured later humid period for eastern Amazonia (based for example on records from the CarajaH s mountains) between 12,400 and 8800 cal yr B.P. (ca. 10,700 and 7700 C yr B.P.) and with an onset at ca. 13,000 C yr B.P. They suggest a somewhat northward shifted summer ITCZ due to massive, astronomical controlled (precession), insolation changes over South America during this period. However, to what extend the coastal area of NE Brazil was a!ected by such ITCZ movements is unclear. Considering more global-scale atmospheric changes, the correspondence of the repeated humid periods over NE Brazil with the North Atlantic Bond cycles (maximum cooling) would rather imply a slightly southward displacement of the ITCZ during the Northern Hemisphere stadials, directly a!ecting the precipitation in the study area. Changes of the highpressure cell over the southern Atlantic were probably also important. Intensi"ed SE trade winds and surface currents during each of the humid periods (Little et al., 1997; Arz et al., 1998) could have induced a stronger moisture supply from the open Atlantic Ocean thus favoring orographic rains in the coastal area of NE Brazil, a fact which is also observed on modern seasonal scale (Rao et al., 1993). Furthermore, a strong northward shift of Antarctic cold fronts over the eastern Brazilian highland to NE Brazil, could explain the higher precipitation rates and a markedly shorter annual dry season during the Lateglacial. The occurrence of a wet period during the early Holocene (8800 to 7500 C yr BP) in southeastern Brazil (Lago do Pires record), may be the result that polar cold fronts reached only so far north at that time. The retreat of the Antarctic cold fronts from NE Brazil to southeastern Brazil at the end of the Lateglacial may explain the return to semi-arid conditions. The same mechanism was suggested by Martin et al. (1993) for South American climate changes during the last 7000 yr, which they interpreted to be related to long-term El Nin o H. Behling et al. / Quaternary Science Reviews 19 (2000) 981}994 activity. Questionable, is whether Antarctic cold fronts could reach a latitude of ca. 43S, because cold fronts penetrating as far north as the coastal areas of Brazil would expect to rather be blocked by SW trade winds. 6. Conclusion Pollen analysis of modern surface samples and the well dated marine core GeoB 3104-1 located o! NE Brazilian continental slope lead to conclusions that contribute to the understanding of vegetational and climatic changes on the adjacent continent. Pollen data from river and lake surface sediments and forest soil surface samples provide a modern pollen analog of NE Brazilian vegetation types. Di!erent herb taxa are predominant, re#ecting primarily open caatinga vegetation types. Marine pollen data indicate the occurrence of caatinga vegetation in NE Brazil during the recorded part of the Last Glacial and early Holocene period (42,000 } 8500 C yr BP), re#ecting most of the time semi-arid conditions. The increase of transport of terrigenous material to the Atlantic Ocean, which contain high concentration of pollen and especially fern spores during relatively short intervals around 40,000, 33,000 and 24,000 C yr BP, indicate periods of higher precipitation. The wettest climate is found from 15,500 to 11,800 C yr BP. These climatic changes can be explained by variations in the tropical/subtropical wind "eld. Timing, duration, and amplitude of these climatic events are comparable with the Daansgaard-Oeschger and Bond cycles, well known from higher North Atlantic high latitudes, suggesting the presence of a teleconnection in the climate system. Only the Lateglacial period allowed an expansion of mountain forests, #oodplain forest and gallery forests along rivers, suggesting cold and very wet climatic conditions with short annual dry seasons. Floristic exchanges between Atlantic rain forest and Amazon rain forest vice versa was during the Lateglacial period possible. The consistent annual movement of the ITCZ over NE Brazil, the strong in#uence of the Antarctic cold fronts and changes of the high pressure cell over the southern Atlantic may explain the very wet Lateglacial period in NE Brazil. Acknowledgements The authors thank the RV Victor Hensen cruise JOPS II members for the collection of marine sediment cores. This cruise was supported by the Federal Department for Science and Technology in Germany, and the Department of Environment, Brazil. Dr. Matthias Tintelnot, from the Senckenberg Institute in Wilhelmshaven (Germany) is thanked for some river surface samples from NE Brazil. 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