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JOURNAL OF QUATERNARY SCIENCE (2004) 19(1) 3547 Copyright 2004 John Wiley & Sons, Ltd.

. Published online in Wiley InterScience ( DOI: 10.1002/jqs.813

Millennial-scale climate variability in northwest Patagonia over the last 15 000 yr

PATRICIO I. MORENO* gicas, Facultad de Ciencias, Universidad de Chile, Casilla 653, Santiago, Chile Departamento de Ciencias Ecolo
Moreno, P. I. 2004. Millennial-scale climate variability in northwest Patagonia over the last 15 000 yr. Vol. 19 pp. 3547. ISSN 0267-8179. Received 8 October 2002; Revised 5 October 2003; Accepted 12 October 2003

ABSTRACT: A pollen record from Lago Condorito (41  45S, 73  07W) shows prominent vegetation and climate changes at millennial time-scales, superimposed on multimillennial trends in temperature and westerly activity in northwest Patagonia during the past 15 000 yr. The record shows that evergreen temperate rainforests have dominated the landscape over this interval, with oristic changes ranging from cold-resistant North Patagonian forests with podocarp conifers to Valdivian forests with thermophilous, summer-drought resistant species. The long-term trend shows that cool-temperate and humid conditions prevailed between 15 000 and 11 000 cal. yr BP, followed by an extreme warm and dry phase between 11 000 and 7600 cal. yr BP, and subsequent cooling events and increase in precipitation that peaked at ca. 5000 cal. yr BP, when Southern Hemisphere alpine glaciers achieved their rst Neoglacial maximum. Modern conditions were established at ca. 1800 cal. yr BP, following a warm and dry phase between ca. 2900 and 1800 cal. yr BP. These results suggest that millennial-scale climate variability during deglacial and post-glacial times also affected the mid-latitude region of the South Pacic, supporting the idea that changes in the tropical Pacic might be a key factor in the initiation and/or propagation of millennial-scale climate variability at regional, hemispheric and global scales. Copyright 2004 John Wiley & Sons, Ltd.
KEYWORDS: millennial-scale climate variability; Patagonia; temperate rainforests; pollen; Holocene; Late-glacial.

High-resolution palaeoclimate records from Greenland, Antarctica (Stauffer et al., 1998), the low-latitude Pacic Ocean (Behl and Kennett, 1996), the Arabian Sea (Leuschner and Sirocko, 2000) and the North Atlantic Ocean (Bond and Lotti, 1995; Bond et al., 1997) were rst to demonstrate the occurrence of (quasi-) cyclic, millennial-scale climate uctuations during and since the last ice age. Ever since then, continental records have conrmed this pattern (Allen et al., 1999; Baker et al., 2001; Mayle et al., 2000; Pisias et al., 2001; nchez Gon i et al., 2002) indicating that millennial-scale Sa variability is a widespread and dominant feature of the climate system. However, the mechanisms driving these uctuations at hemispheric and global scales remain uncertain. Although the exact phasing and geographical extent of these events is imperfectly known, it has been suggested that they originated from reorganisations of the coupled oceanatmosphere system in the North Atlantic (Alley et al., 1999), the tropical Pacic (Cane and Clement, 1999) or Southern Oceans (Ninnemann et al., 1999), and were transmitted elsewhere through atmospheric

gicas, * Correspondence to: Prof. P. I. Moreno, Departamento de Ciencias Ecolo Facultad de Ciencias, Universidad de Chile, Casilla 653, Santiago, Chile. E-mail:

and/or oceanic teleconnections. Understanding the causes and consequences of millennial time-scale variability requires knowledge of their extent and chronology, especially in areas where climatic controls (e. g. presence/absence of deep-ocean ventilation and eastern boundary currents, easterly versus westerly zonal ow, low versus high-latitude regions, Antarctic/ Arctic regions) are most contrasting. Additional high-resolution, precisely dated land and marine records from the poorly understood South Pacic region may help clarify the contribution of low- and high-latitude climate processes in hemispheric and global climate changes. Few palaeoclimate records in this region have examined in detail the timing, frequency and character of palaeoclimate events during the past 15 000 yr. These data are crucial in identifying the extent and interhemispheric phasing of millennial-scale variability, and understanding the climatic mechanisms that led to extreme and contrasting phases within the Holocene, such as the climatic optimum/Neoglaciation, along with the onset of high-frequency climate variability, including the birth oSouthern Oscillation phenomenon. of the El Nin This study presents a high-resolution, precisely dated pollen record that spans the past 15 000 yr. These data allow assessment of the following questions. (i) What was the timing and direction of vegetation changes in northwest Patagonia over the past 15 000 yr? (ii) Did vegetation changes in the study area occur in a gradual or abrupt manner? (iii) How did the southern westerlies vary during deglacial and postglacial times? (iv) Is



there evidence for a millennial- (and multimillennial) scale pulsing in vegetation and climate changes in mid-latitude South America? (v) If so, what is the phasing relationship with palaeoclimate records from polar latitudes (Antarctica, Greenland)? (vi) Has re played a signicant role in the development of rainforest vegetation since the end of the last ice age?

extensive European settlement started in 1850. This process disturbed the native vegetation in the Central Depression between the cities of Valdivia and Puerto Montt, leading to a rapid change from a landscape dominated by native forests to one dominated by agriculture and pasture land with small fragmented patches of forests. Most of this conversion was achieved by the end of the nineteenth century (Fraver et al., 1999; Lara et al., 1999).

Study area
Northwest Patagonia (40  43  S) is ideal for the study of past vegetation and climate changes, thanks to the presence of temperate rainforests highly sensitive to climatic gradients, the abundance of sites with high sediment accumulation rates and continuous deposition since the last glaciation, and the feasibility of studying variations of the southern westerlies during deglacial and post-glacial times. Abundant precipitation in this region, brought by westerly winds, occurs evenly throughout the year, with decreased frontal activity during the summer months. Seasonal shifts in the latitudinal position of the westerly storm tracks, along with varying intensity of coastal upwelling and the Humboldt Current, result from meridional sea-surface temperature gradients and the interaction between the subtropical Pacic high-pressure cell and the polar lowpressure belt (Miller, 1976). Montecinos et al. (2000) found a o signicant tendency for relatively dry summers during El Nin events in the area between 38  and 41  S. An anomaly of oppo a events, indicating that intersite sign is evident during La Nin annual-scale variability also exerts an important inuence on precipitation regimes in the study area. Temperate rainforests form the predominant vegetation in the Chilean Lake District between 0 and 1200 m elevation in the mountain ranges at 41  S. Strong climatic gradients lead to zonation of forest communities along altitudinal and latitudinal gradients, providing a modern analogue for interpreting n, 1980; Pa ez palynological records (Heusser, 1974; Villagra et al., 1996). Modern analyses of vegetation and pollen rain ez et al., 1996) revealed three everon the Andes at 42  S (Pa green forest types along a transect of increasing altitude. Zonation of these communities reects increasing amounts of precipitation, and declines in mean annual temperatures and precipitation seasonality. Although these forest communities share many taxa, the presence of key indicator species in the modern pollen rain allow recognition of: (i) Valdivian rainforest with the conspicuous trees Eucryphia cordifolia and Aextoxicon punctatum; (ii) North Patagonian rainforest dominated by species of the genus Nothofagus, species of the family Myrtaceae, many other evergreen trees, vines and epiphytes, but lacking conifers; and (iii) North Patagonian rainforest with podocarpaceous (Podocarpus nubigena, Saxegothaea conspicua) and cupressaceous conifers (Fitzroya cupressoides, Pilgerodendron uviferum). Although strong gradients in temperature, precipitation and rainfall seasonality lead to latitudinal and altitudinal segregation of forest communities, geographical overlap does take place between Valdivian and North Patagonian rainforests at smaller spatial scales. Fine-scale mosaics are present in the , in lowlands of the Lake District and Isla Grande de Chiloe what could be considered a climatic transition or tension zone for hygrophilous, cold-resistant North Patagonian trees, in particular conifers (Podocarpus nubigena, Saxegothaea conspicua and Fitzroya cupressoides). Large-scale disturbance by human-set res for land-clearing for agriculture and pasture land, grazing and logging following
Copyright 2004 John Wiley & Sons, Ltd.

Materials and methods

This study is based on a sediment core from Lago Condorito (41  45S, 73  07W), a small elliptical pond located in the lowlands of the southern Chilean Lake District (Fig. 1). Lago Condorito lies within a closed basin with a single concave depression, and a maximum water depth of 270 cm. The lake is situated in an intermorainal depression atop a Llanquihueage moraine belt that surrounds the western rim of the Seno seaway, near the city of Puerto Montt (Denton Reloncav et al., 1999). Sediment cores for this study were collected with a 5-cmdiameter Wright square-rod piston corer. Three overlapping sediment cores were obtained from the centre of Lago Condorito to develop high-resolution, continuous pollen and charcoal records devoid of hiatuses. The stratigraphy of the cores was documented with the aid of X-radiographs, along with textural description of the sediments. The organic and carbonate content of the sediments was determined by loss-on-ignition at 550  C for 2 h, and 925  C for 4 h, after drying overnight at 70  C (modied from Bengtsson and Enell (1986) and Dean (1974)). A radiocarbon chronology was developed based on accelerator mass spectrometry (AMS) radiocarbon dates obtained from bulk lake sediment and plant macrofossils of terrestrial origin, along with conventional dates on bulk lake sediment, considering the absence of hard water and reservoir effects throughout the region. Samples for pollen analysis were processed following standard procedures, which include KOH, sieving (<120 mm), deoculation, HF and acetolysis (Faegri and Iversen, 1989). Tablets with exotic Lycopodium spores were added to allow calculation of concentration and accumulation rates (inux abundance cm2 yr1) for pollen, spores and microscopic charcoal particles (Stokmarr, 1971). A total of 300 pollen grains including upland trees, shrubs and herbs were counted for each level (terrestrial pollen basic sum). The abundance of paludal/aquatic taxa, as well as pteridophytes were added to separate sums, and their percentage values were calculated with reference to the supersums total pollen (terrestrial pollen paludal/aquatics) and total pollen and spores (total pollen pteridophytes). This treatment allows a better appreciation of changes in the upland vascular vegetation, separating its variation from site or substrate-specic signals on the fern/aquatic vegetation. A paleovegetation index was calculated based on the percentage abundance of the palynomorph Eucryphia/ Caldcluvia 1 divided by the combined abundance of the species Podocarpus nubigena and Saxegothaea conspicua 1. The resulting values were then normalised following conversion to the logarithmic scale. The rationale behind this index lies in the evident segregation of these taxa along broad-scale geographical and climatic gradients, and their excellent indicator value of rainforest communities with contrasting oristic assemblages. The Eucryphia/Caldcluvia to Podocarpaceae
J. Quaternary Sci., Vol. 19(1) 3547 (2004)



. Right panel: detail of the surface drainage system of the Figure 1 Left panel: map of the Chilean Lake District and adjacent Isla Grande de Chiloe study area

ratio constitutes a useful parameter to quantify: (i) the degree of mixture of Valdivian and North Patagonian rainforest communities; and (ii) the relative position of a fossil pollen sample along a gradient ranging from a warm/temperate, seasonally dry climate characteristic of the lowland environments between 39  and 43  S, to a cool-temperate condition with higher amounts of precipitation evenly distributed throughout the year, characteristic of the mid- to high elevations of the Coastal and Andean Ranges at 40  43  S and the maritime region of the Chilean channels between 43  48  S. Pollen analysis was performed at 400 and 1000 magnication on a Nikon Labophot microscope (Late-glacial and early Holocene portions) and a Zeiss Axioscope microscope (mid- and late Holocene portions). Pollen assemblages were dened based on a stratigraphically constrained CONISS ordination, using square-root transformed data and the Edward and Cavalli-Sforzas chord distance as a dissimilarity coefcient. The rates-of-change parameter was calculated to quantify the magnitude/rapidity of vegetation changes over the past 15 000 yr. This was done by smoothing the pollen percentage data with a ve-point moving average, and interpolating pollen samples at regular 150-yr intervals. Both the CONISS ordination and rates-of-change analysis were performed on all terrestrial pollen taxa (excluding pteridophytes) that reach percentages !2%, after recalculating sums and percentages (Jacobson et al., 1987). Microscopic charcoal particles were counted individually in each pollen slide to reconstruct the occurrence of past res in the landscape. Rather than specifying a specic size, severity, magnitude or number of individual re events, microscopic charcoal provides a generalised description of past re occurrence, typically ranging from decades to centuries or more. Charcoal abundance is expressed as inux, considering the potential bias introduced by varying sediment accumulation rates.
Copyright 2004 John Wiley & Sons, Ltd.

Stratigraphy and chronology
All analyses were conducted on a sedimentary core 930 cm long (core PM10, Fig. 2) retrieved from the deepest part of the basin, considering its stratigraphical range and completeness. The sediments of core PM10 consist largely of gyttja, silty/sandy gyttja between 604 and 692 cm and 120 and 162 cm, and a basal peat layer. Two prominent volcanic ash layers are evident between 699 and 726 (ca. 11 000 cal. yr BP) and 6670 cm depth (ca. 130 cal. yr BP). The carbonate content of the sediments is negligible or nil (<4%). A total of 18 radiocarbon dates afford chronological control for the Lago Condorito record (Table 1). Radiocarbon dates were converted to calendar years BP using the Calib 4.1.2 program (Stuiver et al., 1998). The date of sample A-6661 was excluded from the age model because it is too young and should be regarded as a minimum age for the onset of deposition in Lago Condorito (Moreno, 2000). Based on the radiocarbon and calendar dates two age models were developed: fth-order polynomial regressions on the radiocarbon dates (R2 0.999, P < 0.0001) and calibrated radiocarbon dates (R2 0.997 and P < 0.0001) (Fig. 2). The age model based on calibrated radiocarbon dates was chosen to assign interpolated ages to the palynological results presented in this paper.

Palynological results
The pollen, spore, and charcoal content of 260 samples were analysed at an average time resolution of ca. 60 yr between
J. Quaternary Sci., Vol. 19(1) 3547 (2004)



Figure 2 Stratigraphical column of core PM10 showing all 14C dates and various stratigraphical parameters. Dashed lines mark the onset of major events in the pollen stratigraphy (see Figs 3 and 4 and Table 2). The right portion of the gure shows the radiocarbon and calendar age models Table 1 Radiocarbon dates from core PM10. Radiocarbon-age errors correspond to 1 NSRL, Laboratory for AMS Radiocarbon Preparation and Research, University of Colorado INSTAAR; A, University of Arizona Geochemistry Laboratory; Beta, Beta Analytic Depth (cm) 6868.5 109111 180180.5 230231 305306 370371 410411 437439 510511 575578 613615 685688 710713 821824 848852 863873 873886 928930 Material dated Wood Wood Bulk Bulk Bulk Bulk Bulk Leaf Bulk Bulk Bulk Bulk Bulk Bulk Bulk Bulk Bulk Bulk

C year BP 1 error 90 35 1680 80 2750 45 3710 90 4370 50 5190 50 5570 50 6020 45 7620 130 8570 45 9110 45 9680 85 10 060 60 11 265 65 11 435 80 12 230 140 12 170 90 12 330 130

13C 26.7 30.4 27.9 28.5 28.6 28.9 29.1 26.6 29.3 29.3 30.1 30.5 28.4 29.9 29.5 30.0 32.1 29.0

Calendar year BP (1 range) (60, 42, 0) 0 1694 (1562) 1519 2917 (2848) 2780 4221 (4084, 4028, 4005) 3911 5030 (4954, 4951, 4872) 4860 5988 (5928) 5910 6406 (6388, 6372, 6312) 6297 6894 (6858, 6821, 6804) 6757 8540 (8405) 8335 9548 (9535) 9528 10 071 (10 035) 10 006 10 976 (10 936) 10 611 11 825 (11 513, 11 361, 11 339, 11 273, 11 266, 11 229) 11 089 13 263 (13 172) 13 091 13 465 (13 347) 13 245 14 515 (14 278) 14 066 14 381 (14 202) 14 040 14 648 (14 411) 14 198

Laboratory reference NSRL-10724 NSRL-10723 NSRL-10722 NSRL-11076 NSRL-11077 NSRL-11078 NSRL-11079 NSRL-10725 NSRL-11080 NSRL-10721 A-8587 A-8070 A-8069 A-8068 A-8067 Beta-60352 A-8066 A-6661

levels (Figs 3 and 4). A stratigraphically constrained CONISS ordination revealed 13 pollen assemblage zones, which highlight the major patterns in the variation of key indicator taxa (Fig. 3 and Table 2). Pollen zones were named considering a hierarchy of the three or four most abundant taxa per pollen zone (mean values) (Table 2). A palaeoclimate event number was assigned to each transition between pollen assemblage zones in descending order toward the present to facilitate discussion in the nal part of the paper, and for establishing comparisons with other palaeoclimate records. Here follows a description of the pollen assemblage zones. The mean percentage abundance of each taxon is expressed in parentheses whenever pertinent.
Copyright 2004 John Wiley & Sons, Ltd.

Pollen zone CO-1 (15 00013 800 cal. yr BP) comprises the assemblage MyrtaceaeNothofagus dombeyi-typeFitzroya/ Pilgerodendron, along with Pseudopanax laetevirens (6.4%), Gramineae (6.2%), Hydrangea (4.7%), Lomatia/Gevuina (3.2%), Escallonia (2.6%), the ferns Blechnum-type (4.1%), Hymenophyllaceae (2%), and traces (mean < 2%) of Podocarpus nubigena, Umbelliferae, Donatia fascicularis, Astelia pumila and Polypodium feullei. Peak abundance of Gramineae in the basal part of the record correlates with charcoal peaks between ca. 15 00014 500 cal. yr BP. Pollen zone CO-2a (event 12, 13 80012 500 cal. yr BP) features the dominance of Nothofagus dombeyi-typeMyrtaceae Podocarpus nubigena, accompanied by Pseudopanax laeteviJ. Quaternary Sci., Vol. 19(1) 3547 (2004)



Figure 3 Percentage diagram of selected taxa from core PM10. Dashed lines mark the onset of major events in the pollen stratigraphy

Copyright 2004 John Wiley & Sons, Ltd.

J. Quaternary Sci., Vol. 19(1) 3547 (2004)



rens (15%), Hydrangea (14.3%), Weinmannia trichosperma (3.1%), Blechnum-type (2.7%), Hymenophyllaceae (2.6%), Escallonia (2.1%), and traces of Fitzroya/Pilgerodendron, Lomatia/Gevuina, Gramineae, Donatia fascicularis, Astelia pumila and Polypodium feullei. The accumulation-rate diagram shows that the expansion of Nothofagus dombeyi-type and Podocarpus nubigena at 13 80012 500 cal. yr BP was an abrupt event. An isolated charcoal peak is evident at 13 400 cal. yr BP. Pollen zone CO-2b (event 11, 12 50011 000 cal. yr BP) comprises the assemblage Weinmannia trichospermaNothofagus dombeyi-typeHydrangea, with prominent declines in

Myrtaceae (from 18.6% to 6.2%), Podocarpus nubigena (from 15.4% to 6%), Pseudopanax laetevirens (from 15% to 5.1%), and increases in Tepualia stipularis (from a near absence to 2.7%) and Gramineae (from 1.8% to 4.2%). Other taxa include Escallonia (2.4%), Blechnum-type (2.1%), and traces of Lomatia/Gevuina, Donatia fascicularis, Astelia pumila, Hymenophyllaceae and Polypodium feullei. The accumulation-rate diagram shows that the expansions of Weinmannia trichosperma and Tepualia stipularis at 12 500 cal. yr BP and charcoal were abrupt. Charcoal reached abundance maxima at 12 50012 000 cal. yr BP and 11 40011 000 cal. yr BP.

Figure 4 Inux diagrams of selected taxa from core PM10. Calculations are based on the calendar age scale to avoid biases introduced by the radiocarboncalendar age differential. Inux scales among taxa vary for visual depiction only. Dashed lines mark the onset of major events in the pollen stratigraphy
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Figure 4 (Continued ) Table 2 Summary of the pollen zones from Lago Condorito, indicating their age range and dominant taxa Age range (1000 cal. yr BP) 01.8 1.82.9 2.94.1 4.14.9 4.95.7 5.76.9 6.97.6 7.69.3 9.310 1011 1112.5 12.513.8 13.815 Pollen zone CO-7b CO-7a CO-6c CO-6b CO-6a CO-5b CO-5a CO-4b CO-4a CO-3 CO-2b CO-2a CO-1 Dominant taxa

Weinmannia trichospermaNothofagus dombeyi-typeTepualia stipularis Nothofagus dombeyi-typeWeinmannia trichospermaTepualia stipularis Nothofagus dombeyi-typeSaxegothaea conspicuaWeinmannia trichospermaEucryphia cordifolia/ Caldcluvia paniculata Nothofagus dombeyi-typeSaxegothaea conspicuaWeinmannia trichospermaTepualia stipularis Nothofagus dombeyi-typeSaxegothaea conspicuaTepualia stipularisWeinmannia trichosperma Nothofagus dombeyi-typeTepualia stipularisSaxegothaea conspicua Tepualia stipularisNothofagus dombeyi-typeEucryphia cordifolia/Caldcluvia paniculataGramineae Tepualia stipularisEucryphia cordifolia/Caldcluvia paniculataNothofagus dombeyi-type Eucryphia cordifolia/Caldcluvia paniculataWeinmannia trichospermaGramineae Weinmannia trichospermaTepualia stipularisMyrtaceae Weinmannia trichospermaNothofagus dombeyi-typeHydrangea Nothofagus dombeyi-typeMyrtaceaePodocarpus nubigena MyrtaceaeNothofagus dombeyi-typeFitzroya/Pilgerodendron

Pollen zone CO-3 (event 10, 11 00010 000 cal. yr BP) is characterised by the assemblage Weinmannia trichosperma Tepualia stipularisMyrtaceae, along with increases in Gramineae (9.8%), Pseudopanax laetevirens (8.6%), Hydrangea (from 6.7% to 8.1%), Escallonia (7.7%), Lomatia/Gevuina (5.8%) and traces of Eucryphia/Caldcluvia. Other taxa decline (Nothofagus dombeyi-type, from 15% to 8%) or disappear
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(Podocarpus nubigena, Donatia fascicularis and Astelia pumila). Charcoal accumulation rates peak during this zone, reaching the highest abundance of the past 15 000 yr. Pollen zone CO-4a (event 9, 10 0009300 cal. yr BP) is dominated by the assemblage Eucryphia/Caldcluvia Weinmannia trichospermaGramineae. These taxa are associated with other trees such as Tepualia stipularis (9.6%),
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Pseudopanax laetevirens (7.2%), Nothofagus dombeyi-type (6.5%), Lomatia/Gevuina (4.9%), Myrtaceae (4.6%) and the vine Hydrangea (5%). Charcoal declines, stabilizing at intermediate values. Pollen zone CO-4b (event 8, 93007600 cal. yr BP) features the dominance of Tepualia stipularis, Eucryphia/Caldcluvia and Nothofagus dombeyi-type, along with Hydrangea (6.9%), Myrtaceae (4.2%), and declining Gramineae (from 10% to 6.7%), Weinmannia trichosperma (from 15.2% to 5.8%) and traces of Saxegothaea conspicua. The most salient aspect of this pollen zone is the abrupt increase in Tepualia stipularis both in per cent (from 9.6% to 30.9%) and accumulation rate abundance, along with the persistence of Eucryphia cordifolia/Caldcluvia paniculata in a high-stand plateau (28.6%). Charcoal accumulation rates reached a peak at ca. 8800 cal. yr BP and declined thereafter to minimum values. Pollen zone CO-5a (event 7, 76006900 cal. yr BP) comprises the assemblage Tepualia stipularisNothofagus dombeyitypeEucryphia cordifolia/Caldcluvia paniculataGramineae, which is accompanied by Hydrangea (7.7%), Weinmannia trichosperma (7.3%), Myrtaceae (4.7%), Saxegothaea conspicua (2.8%), and traces of Podocarpus nubigena, Pseudopanax laetevirens and Drimys. The accumulation-rate diagram shows that the decline of Eucryphia cordifolia/Caldcluvia paniculata and Tepualia stipularis at 7600 cal. yr BP was abrupt, and reveals that this pollen zone is transitional between CO-4b and CO-5b. Pollen zone CO-5b (event 6, 69005700 cal. yr BP) is dominated by Nothofagus dombeyi-type, Tepualia stipularis and Saxegothaea conspicua, and includes Weinmannia trichosperma (8.4%), Gramineae (6.2%), Myrtaceae (6.1%), Hydrangea (5.4%) and Eucryphia cordifolia/Caldcluvia paniculata (3.2%), along with traces of Podocarpus nubigena, Pseudopanax laetevirens and Drimys. Pollen zone CO-6a (event 5, 57004900 cal. yr BP) features the dominance of Nothofagus dombeyi-typeSaxegothaea conspicuaTepualia stipularis, along with Weinmannia trichosperma (8.1%), Hydrangea (5.5%), Gramineae (4.2%), Lomatia/Gevuina (3.5%), Eucryphia/Caldcluvia (2.8%) and Podocarpus nubigena (2.1%). The accumulation-rate diagram shows an abrupt increase in Nothofagus dombeyi-type and Saxegothaea conspicua. Pollen zone CO-6b (event 4, 49004100 cal. yr BP) is characterised by the assemblage Nothofagus dombeyi-type Saxegothaea conspicuaWeinmannia trichosperma. Other taxa include Tepualia stipularis (8.5%), Hydrangea (6.5%), Eucryphia cordifolia/Caldcluvia paniculata (4.4%), Gramineae (2.9%), Lomatia/Gevuina (2.2%) and traces of Podocarpus nubigena. The accumulation-rate diagram shows that Nothofagus dombeyi-type and Saxegothaea conspicua reached their peak Holocene abundance between 5200 and 4500 cal. yr BP. Pollen zone CO-6c (event 3, 41002900 cal. yr BP) features the dominance of the same three main taxa as above, and shows a two-step increase in Eucryphia cordifolia/Caldcluvia paniculata: an abrupt jump between 4100 and 3800 cal. yr BP (from 4.4% to 16.7%), and a subsequent gradual and highly variable rising trend that culminated at 1800 cal. yr BP. Charcoal accumulation rates rose slightly during this zone. Pollen zone CO-7a (event 2, 29001800 cal. yr BP) with the dominant Nothofagus dombeyi-typeWeinmannia trichosperma Saxegothaea conspicua assemblage, followed in abundance by Tepualia stipularis (10.1%), Eucryphia cordifolia/Caldcluvia paniculata (9.2%), Hydrangea (3.9%), Pseudopanax laetevirens (2.7%), Podocarpus nubigena (2.5%), Gramineae (2.3%), Lomatia/Gevuina (2.2%) and traces of Myrtaceae. Pollen zone CO-7b (event 1, 18000 cal. yr BP) is characterised by the assemblage Weinmannia trichosperma
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Nothofagus dombeyi-typeTepualia stipularis. Other important taxa are Eucryphia cordifolia/Caldcluvia paniculata (9.7%), Saxegothaea conspicua (6.2%), Pseudopanax laetevirens (5.3%), Hydrangea (3.6%), Lomatia/Gevuina (3.1%) and Gramineae (2.2%), along with traces of Podocarpus nubigena and Myrtaceae.

Environmental reconstruction
Stratigraphy and chronology
The X-radiographs, loss-on-ignition, lithostratigraphy and radiocarbon chronology suggest undisturbed, continuous deposition in the Lago Condorito basin during the past 15 000 yr. Peat accumulation between 15 000 and 13 800 cal. yr BP suggests a shallow environment peripheral to a small pond. The onset of gyttja deposition at 13 800 cal. yr BP and its subsequent persistence suggests a rise in water table and the establishment of a perennial lake. Gradual transitions between gyttja to sandy or silty gyttja (between 699 and 621 and 178 and 110 cm, equivalent to ca. 11 00010 000 and ca. 3000 2000 cal. yr BP, respectively) are expressed as decreases in the percentage organic content (decline in the concentration of organic content and a rise in the concentration of inorganic content) (Fig. 2). These inorganic-enriched zones may represent internal reworking of clastic material in the Lago Condorito basin resulting from the resuspension of volcanic ash and/or falls in lake-level. Aeolian transport of silt- or sand-sized particles is unlikely, considering that: (i) dense rainforests persisted during these intervals (arboreal pollen with a mean of 87 and 97%, respectively) and thus may have prevented any signicant entrainment and deposition of wind-blown ne-grained sediment; (ii) Lago Condorito is a closed-basin pond, with no possible input of allochtonous sediments via uvial processes; and (iii) the silty gyttja horizon between 178 and 110 cm depth is unrelated to the deposition of volcanic ejecta. Furthermore, these inorganic-enriched layers occur at times when the local vegetation changed toward relatively drier conditions (see discussion below).

Pollen and charcoal records

The palynological results from Lago Condorito suggest that temperate evergreen rainforests have dominated the lowlands of the Chilean Lake District over the past 15 000 yr (arboreal pollen: 93.7%). A highly dynamic picture of vegetation change emerges from the Lago Condorito record, as revealed by the pollen assemblage zones (n 13). Given the abruptness and character of vegetation change (see discussion below) across pollen zone boundaries, these transitions are interpreted as being driven by climate change. Consequently, each boundary will be treated as a palaeoclimate event. The record from core PM10 shows the presence of North Patagonian rainforest taxa between ca. 15 000 and 13 800 cal. yr BP, followed by the expansion of Nothofagus dombeyi-type and the conifer Podocarpus nubigena, along with a prominent decline in other trees between 13 800 and 11 000 cal. yr BP (Fig. 5). Podocarpaceae pollen (P. nubigena, Saxegothaea conspicua) is most abundant today in surface ez et al., samples from mid- to high-elevation rainforests (Pa 1996), hence the pollen record between 13 800 and 11 000 cal. yr BP suggests a lowering of treeline under cooltemperate and humid conditions (event 12). This inference
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Figure 5 Summary pollen diagram and results of the rates-of-change analysis on the Lago Condorito record. Dashed lines mark the onset of major events in the pollen stratigraphy. The numbers on the right-hand side of the diagram indicate the palaeoclimate events discussed in the text

is supported by the establishment of a deeper perennial lake, as evidenced by the sequence from peat sediments to gyttja at 13 800 cal. yr BP. An abrupt rise of the shade-intolerant trees Weinmannia trichosperma and Tepualia stipularis between 12 500 and 11 000 cal. yr BP (event 11) is superimposed on this trend, coeval with major increases in charcoal particles. A decline in Nothofagus dombeyi-type, Podocarpus nubigena, Weinmannia trichosperma and Tepualia stipularis occurred at 11 000 cal. yr BP (event 10), along with a major expansion of other temperate rainforest taxa between 11 000 and 10 000 cal. yr BP. These data indicate a decline in coldtolerant taxa and expansion of other trees with broad distributions in geographical and climate space, suggesting a warming event at 11 000 cal. yr BP and onset of the Holocene. A rapid expansion of Eucryphia cordifolia/Caldcluvia paniculata occurred at 10 000 cal. yr BP (event 9), followed by a marked increase in T. stipularis by 9300 cal. yr BP (event 8). Both taxa attained high abundance until 7600 cal. yr BP, after which they plummeted to their minima (event 7). The dominance of Eucryphia cordifolia/Caldcluvia paniculata during the early Holocene suggests the establishment of warmer conditions, decreased mean annual precipitation and, quite possibly, increase in rainfall seasonality. The increase of T. stipularis at 9300 cal. yr BP might represent an increase in humidity, as suggested by a near-synchronous drop in charcoal particles, and its occurrence in waterlogged environments and in the temperate, hyperhumid regions of southern Chile. The decline of Eucryphia cordifolia/Caldcluvia paniculata at 6900 cal. yr BP (event 6) attests to an abrupt end of the early Holocene climatic optimum, most probably triggered by cooler conditions and an increase in precipitation. Fire occurrence near Lago Condorito has remained low or absent since the last peak at 8800 cal. yr BP. Nothofagus dombeyi-type, Saxegothaea conspicua and Podocarpus nubigena expanded in pulses centred at 6900 and 5700 cal. yr BP, suggesting the onset (event 6) and subsequent intensication (event 5) of cooler conditions. This trend, which culminated at ca. 4500 cal. yr BP, was followed by a decline in S. conspicua (event 4), an abrupt expansion of Eucryphia cordifolia/Caldcluvia paniculata between 4100 and 3900 cal. yr BP and a subsequent re-expansion of S. conspicua that started at 3900 cal. yr BP (event 3). The percentage and inux data show high variability between ca. 3500 and 2000 cal. yr BP and the onset of an irreversible trend toward minimum values in Nothofagus dombeyi-type, S. conspicua
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and P. nubigena at ca. 2900 cal. yr BP (event 2). The loss-onignition analysis reveals a drop in the concentration of total organic matter and a concomitant increase in the concentration of total inorganic matter between ca. 2900 and 1800 cal. yr BP (Fig. 2). Because these trends are unrelated to allochtonous sediment supply, they are interpreted as resulting from a lake-level drop caused by a decline in precipitation between ca. 2900 and 1800 cal. yr BP. The percentage abundance of Weinmannia trichosperma increased at ca. 1800 cal. yr BP (event 1) and reached a plateau of ca. 30% that has persisted until today.

Palaeoclimatic implications
The pollen data indicate that climate cooled during Late-glacial time, and led to the expansion of cold-resistant rainforest trees between 13 800 and 11 000 cal. yr BP. Within this cool episode, vegetation changed near Lago Condorito in response to small-scale forest res, which Moreno (2000) interpreted as high climate variability and local burning between 12 500 and 11 000 cal. yr BP. A warming trend followed, leading to the expansion of lowland thermophilous trees in pulses centred at 11 000 and 10 000 cal. yr BP. This trend culminated between 10 000 and 7600 cal. yr BP with the warmest conditions of the last glacialinterglacial cycle in this area, judging from long pollen records spanning the early to late stages of the Llanquihue glaciation (Heusser, 1993; Heusser et al., 1999). This early Holocene warm period coincided with a poleward shift of the westerlies into an extreme interglacial mode of westerly n, in press). activity (Moreno and Leo An abrupt reversal in trend characterised the early to midHolocene transition (event 7, Fig. 3), with cooling events at 7600, 6900 and 5700 cal. yr BP. This shift led to the reexpansion of cold-resistant rainforest trees between ca. 6900 and 2900 cal. yr BP, following a transitional period between 7600 and 6900 cal. yr BP. These results strongly suggest the onset of cool-temperate conditions, concomitant with an increase in humidity brought on by an equatorward shift and/ or intensication of the westerlies. The mid-Holocene cooling trend was interrupted by a brief warm and dry event between 4100 and 3800 cal. yr BP. Subsequent warming at ca. 2900 cal. yr BP led to a decline in cold-resistant trees, followed by a rise in precipitation at 1800 cal. yr BP and establishment of modern vegetation and climate.
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The mid-Holocene cooling trend revealed by the Lago Condorito record peaked between 5200 and 4500 cal. yr BP at times when valley glaciers in Patagonia and New Zealand reached their maximum extent during the early Neoglacial advance (Porter, 2000). Because the pollen assemblages during this interval are similar to those during the Late-glacial cool episode, it is possible that both events represent an equivalent amount of temperature depression, i.e. 23  C colder than modern conditions (Heusser et al., 1999; Moreno et al., 1999, 2001; Hajdas et al., 2003). A similar pattern is evident in the 18O record from the Taylor Dome ice-core (Steig et al., 1998) located near the Ross Sea coast of east Antarctica. The nal mid-Holocene cooling pulse (5700 cal. yr BP) in Lago Condorito was nearly synchronous with the onset of o-like variability as inferred from molluscan assemblages El Nin (Sandweiss et al., 1999) and the sedimentary in coastal Peru record of Laguna Aculeo in central Chile (Jenny et al., 2002). This correlation suggests that the onset of ENSO or ENSO-like variability was possible only when the mean climate state of the South Pacic had shifted from an extreme interglacial mode, characteristic of the early Holocene (10 000 7600 cal. yr BP in the Lago Condorito record), to cooler early Neoglacial conditions, which coincide with a mid-Holocene strengthening of the southern westerlies in northwest Patagonia. This conclusion is consistent with the ndings of a weak or non-existent ENSO prior to 7000 cal. yr BP at Laguna Pallcacocha, Ecuador (Moy et al., 2002), less vigorous ENSO in coral records from the tropical Pacic (Tudhope et al., 2001) and diminished ENSO activity in the Ilo region of southern Peru prior to 5300 cal. yr BP (Keefer et al., 2003).

Multimillennial scale variability

Rainforests in northern Patagonia show a high degree of variability over the past 15 000 yr. This variability ranges from Valdivian rainforests with thermophilous species tolerant of occasional summer droughts (Eucryphia cordifolia/Caldcluvia paniculata) to North Patagonian rainforest assemblages with cold-resistant conifers (Podocarpus nubigena and Saxegothaea conspicua). A palaeovegetation index was calculated using these two species with contrasting climatic tolerances (Fig. 6). This index captures the multimillennial trends in the pollen record and allows comparison with other palaeoclimate records and changes in summer insolation at 60  S during the past 15 000 yr. Cool-temperate and humid conditions between 13 800 and 11 500 cal. yr BP (negative values in the palaeovegetation index) correlate with a declining trend that reached minimum values in summer insolation. Warm, relatively dry, conditions between 10 000 and 8000 cal. yr BP (as shown by the preponderance of thermophilous Valdivian elements and, hence, positive departures in the palaeovegetation index) correspond with the beginning of a rising trend in summer insolation. The mid-Holocene cooling trend and increase in precipitation between ca. 7000 and 3000 cal. yr BP (re-expansion of North Patagonian conifers and decline in thermophilous Valdivian trees) was coeval with a sustained increase in summer insolation. Finally, the late Holocene decline in North Patagonian conifers and persistence of thermophilous Valdivian trees (between ca. 3000 and 0 cal. yr BP) correlate with the culmination in the rising trend in summer insolation that had started at ca. 12 000 cal. yr BP. The multimillennial trends in temperature and westerly activity revealed by the Lago Condorito record bear no clear, straightforward, relationship with changes in summer insolation at 60  S (Fig. 6). It is clear that another, still unknown, mechanism must account for the
Copyright 2004 John Wiley & Sons, Ltd.

Figure 6 Comparison between the palaeovegetation index from Lago Condorito (smoothed with a 5-point moving average), iron intensity data from GeoB 33131 (Lamy et al., 2001), the alkenone-based SST reconstruction from core GIK 177482 (Kim et al., 2002), and the December insolation curve for 60  S. The rectangle brackets the early Holocene warm/dry period in the Lago Condorito record

millennial and multimillennial pulsing of palaeoclimate change in northwest Patagonia. The timing and direction of multimillennial trends in temperature and southern westerly activity found in Lago Condorito show a similar pattern to relative sea-surface temperature (SST) trends in the northwest Pacic Ocean (core W8709A-13PC and ODP site 1019) over the past 15 000 yr (Mix et al., 1999). This symmetry suggests a common driving mechanism. Because both regions are affected by coastal upwelling, eastern boundary currents and variations in the strength of subtropical gyre circulation, it is possible that the long-term temperature trends are related to changing equator-to-pole pressure and
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temperature gradients in response to changes in the tropical Pacic region. An apparent discrepancy emerges when comparing the multimillennial trends found in Lago Condorito with marine sedimentary records obtained near the Chilean coast. On one hand, iron intensity data from core GeoB 3313-1 (obtained from the continental slope of the Chilean Lake District at 41  S) were interpreted by Lamy et al. (2001) as variations in westerly activity over the past 7700 cal. yr BP. Based on high iron intensity values between 7700 and 4000 cal. yr BP, Lamy et al. (2001) postulated reduced rainfall at 41  S via a southward shift of the westerlies. On the other hand, an alkenonebased SST reconstruction from marine sediment core GIK 17748-2, obtained from the continental slope of the Valparaiso basin at ca. 33  S, shows that a prominent warming at ca. 7900 cal. yr BP led to a Holocene climate optimum with local SST !18.5  C between ca. 7500 and 5000 cal. yr BP (Kim et al., 2002). These marine-based reconstructions conict with the abrupt vegetation change at 7600 cal. yr BP in the Lago Condorito record, which shows a termination of the Eucryphia cordifolia/Caldcluvia paniculata maximum and the accompanying re-expansion of hygrophilous, cold-resistant North Patagonian conifers and Nothofagus dombeyi-type. The character of this transition clearly indicates an abrupt increase in westerly precipitation and decline in atmospheric temperatures. If replicated by additional studies, this difference could indicate a delayed response of the regional marine record relative to a genuine atmospheric signal underlying the changes in vegetation in the Chilean Lake District.

thermohaline circulation (THC) in the North Atlantic, with the possible exception of a Little Ice Age decline in THC in the Bermuda Rise (Keigwin and Boyle, 2000). A possibility thus exists that millennial-scale cooling during the Holocene, as recorded in Greenland and the North Atlantic Ocean, was a direct consequence of drifting ice itself. If this was the case, a major question remains unsolved: what mechanism triggered marginal uctuations of Northern Hemisphere ice masses during the Holocene? Bond et al. (2001) alluded to a persistent solar inuence on the North Atlantics ca. 1500 yr rhythm during the Holocene. According to this hypothesis, THC variations would act as a magnier and global transmitter of weak solar perturbations via oceanatmosphere feedbacks and teleconnections. A different perspective on millennial-scale variability comes from modelling experiments of insolation-driven changes in ENSO variability during the past 15 000 yr (Clement et al., 2000). Such modelling produced low amplitude and frequency of events during the early Holocene, increasing in both parameters towards the present. Together with past variations in ENSO at millennial and modern-day timescales, as revealed from the Laguna Pallcacocha record (Moy et al., 2003), an emerging picture for a more active role of the tropical Pacic as a trigger for abrupt, globally synchronous climate changes at millennial time-scales is becoming evident (Cane and Clement, 1999; Clement et al., 2000; Pierrehumbert, 2000). The links between processes in the tropical Pacic, North Atlantic ice-rafting, palaeoclimate in the southern mid-latitudes and solar variability are not fully understood, highlighting the need for further modelling and high-resolution palaeoclimate records.

Millennial-scale variability
The Lago Condorito record shows that vegetation and climate changed at millennial time-scales in northwest Patagonia, with prominent events at ca. 1.8, 2.9, 4.1, 4.9, 5.7, 6.9, 7.6, 9.3, 10, 11, 12.5 and 13.8 cal. yr BP. Some of these events are also recorded in previous palynological studies in the study area (Heusser, 1984; Heusser et al., 1999; Moreno et al., 1999; n, in press), thereby adding regional signiMoreno and Leo cance to these results. Additional pollen records from this region, encompassing the onset of extreme glacial climate and subsequent warming during the last termination, show prominent events at 15 800, 17 200, 17 500 and 18 500 cal. yr BP n, in press). Together, (Moreno et al., 1999; Moreno and Leo these results indicate a mean time spacing of 1100 434 yr between climatic events since 20 000 cal. yr BP, falling in the range of millennial-scale climate variability identied in the North Atlantic (Bianchi and McCave, 1999; Bond et al., 1999). Each millennial-scale climate event identied in this study does not follow the same direction and rate of change (Fig. 5) and are in fact superimposed on longer term, multimillennial trends in temperature and westerly activity. This complexity differs from the apparent monotony in the direction of millennialscale climate change in the North Atlantic during the Holocene. Ice rafting and concomitant decreases in SSTs in the subpolar North Atlantic (Bond et al., 1997) correlate with increases in soluble impurities in Holocene ice from Greenland, interpreted as changes in the polar vortex, atmospheric storminess and lowered atmospheric temperatures over Greenland (OBrien et al., 1995). The extent to which these North Atlantic events constitute a driver or a consequence of regional/global climate change is not fully understood. Although millennial-scale changes in the speed of North Atlantic deep-water ow has been reported in the Iceland basin (Bianchi and McCave, 1999), it is clear that freshwater uxes in the North Atlantic during the Holocene were insufcient to trigger radical changes in
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The results from Lago Condorito suggest that millennial-scale climate variability also affected the mid-latitude region of the Southern Hemisphere over the past 15 000 yr. The rates-ofchange analysis reveals that vegetation changed abruptly during most of these millennial-scale events. Vegetation changes at millennial time-scales are superimposed on longer, multimillennial trends in temperature and precipitation. Past changes in precipitation in northwest Patagonia suggest latitudinal shifts in the position and strength of the southern westerlies over the past 15 000 yr. Two cool and humid phases occurred between 13 800 and 11 500 and 7000 and 3000 cal. yr BP, separated by an extreme warm and dry phase during the early Holocene (10 0007600 cal. yr BP). A distinct Late-glacial to early Holocene signal is evident in the charcoal record, with multiple peaks between ca. 15 000 and 14 500 and 12 500 and 8800 cal. yr BP. Fire occurrence near Lago Condorito has remained low or absent since the last peak at 8800 cal. yr BP. Fire played an important role in vegetation changes during the Late-glacial, and possibly during the early Holocene. The results shown in this paper highlight the possible role of the tropical Pacic as a major mover of Holocene climate through its capability of driving: (i) regional-scale changes in SSTs via changes in upwelling intensity (Koutavas et al., 2002), and the Humboldt Current (Sandweiss et al., 2001); (ii) hemispheric-scale inuences by affecting the strength and position of westerly storm tracks via changes in meridional atmospheric pressure gradients (Trenberth, 1991); and (iii) globalscale effects via changes in centres of atmospheric convection (Cane and Clement, 1999; Stott et al., 2002) and overall water vapour content of the tropical atmosphere (Broecker, 1997).
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Acknowledgements I thank G. H. Denton, T. V. Lowell and G. L. Jacobson for their support and assistance during the initial stages of this study, and C. Porter, C. Latorre and T. Henze for logistical support. This research was funded by the US National Science Foundation (Ofce of Climate Dynamics, EPsCOR program, and grant ATM-9809285 to the University of Colorado INSTAAR), NOAA, the National Geographic Society, the Geological Society of America and Fondo Nacional de gico (10 00905, 1030766). This paper co y Tecnolo Desarrollo Cient is a contribution of the Millennium Center for Advanced Studies in Ecology and Research on Biodiversity (grant number P02-051-F-ICM).

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