JOURNAL OF QUATERNARY SCIENCE (2009) 24(3) 248–258 Copyright ß 2008 John Wiley & Sons, Ltd. Published online 27 November 2008 in Wiley InterScience (www.interscience.wiley.com) DOI: 10.1002/jqs.1233 Late Quaternary vegetation and fire history in the northernmost Nothofagus forest region: Mallı́n Vaca Lauquen, Neuquén Province, Argentina VERA MARKGRAF,1,2* CATHY WHITLOCK,3 R. SCOTT ANDERSON2 and ADRIANA GARCÍA4 1 Institute of Arctic and Alpine Research, University of Colorado, Boulder, Colorado, USA 2 Center for Environmental Sciences and Education, and Quaternary Sciences Program, Northern Arizona University, Flagstaff, Arizona, USA 3 Department of Earth Sciences, Montana State University, Bozeman, Montana, USA 4 School of Earth and Environmental Science, University of Wollongong, Wollongong, New South Wales, Australia Markgraf, V., Whitlock, C., Anderson, R. S. and Garcı́a, A. 2009. Late Quaternary vegetation and fire history in the northernmost Nothofagus forest region: Mallı́n Vaca Lauquen, Neuquén Province, Argentina. J. Quaternary Sci., Vol. 24 pp. 248–258. ISSN 0267-8179. Received 28 December 2007; Revised 7 August 2008; Accepted 7 August 2008 ABSTRACT: The last 16 000 cal. a of vegetation, fire and limnological history are described from the steppe-forest ecotone in the northernmost Nothofagus forest region east of the Andes (Mallı́n Vaca Lauquen, Neuquén Province, Argentina, latitude 368 51.3360 S, longitude 718 02.5380 W). Between 16 000 and 14 800 cal. a BP, scrub steppe with substantial open ground expanded in formerly glaciated valleys, whereas Nothofagus–Prumnopitys andina woodland covered mountain slopes. The site was a relatively deep and unproductive small lake at this time. After 14 800 cal. a BP, both steppe and woodland vegetation became denser, indicating increased moisture and temperatures, although not to present levels. The lake was still relatively deep and dystrophic, but became more alkaline by 10 000 cal. a BP. Between 8900 and 5500 cal. a BP, conditions were markedly drier than before; a Cyperaceae marsh developed and disturbance taxa increased. After 5500 cal. a BP, moisture increased but varied greatly, as evidenced by fluctuating water levels and high fire activity from 5500 to 4400 cal. a BP and from 2300 to 1000 cal. a BP. Human activity, in terms of forest clearance and livestock grazing, is documented in the uppermost levels. The evidence of high environmental variability in the middle and late Holocene is consistent with the onset or strengthening of the El Niño– Southern Oscillation, but differences in the timing of fire activity among sites on the west and east sides of the Andes suggest that fuel conditions were important in determining the local occurrence of fire. Copyright # 2008 John Wiley & Sons, Ltd. KEYWORDS: Lateglacial; Holocene vegetation; fire and climate history; mid-latitude Argentine Andes. Introduction The Mallı́n Vaca Lauquen site is a 200 m diameter seasonally inundated wet meadow, on the east side of the Andes of Argentina at the steppe forest ecotone (latitude 368 51.3360 S, longitude 718 02.5380 W, elevation 1567 m). The site was previously the focus of a reconstruction of late Pleistocene/ Holocene vegetation and climate change (Markgraf, 1987). With the more recent focus on the role of past fire activity in shaping structure and distribution of temperate forests in Patagonia and fire-producing climate conditions (Whitlock et al., 2007), re-evaluation of this site, including new pollen, charcoal and plant macrofossil analyses, seemed warranted. Research on documentary evidence, fire-scarred tree rings and sedimentary charcoal records has shown the importance of * Correspondence to: V. Markgraf, 763 N. Pine Cliff Dr., Flagstaff, AZ 86001, USA. E-mail: [email protected] past fires in Patagonia (Veblen et al., 1999; Huber and Markgraf, 2003; Kitzberger and Veblen, 2003; Veblen et al., 2003; Whitlock et al., 2006, 2007). These records document latitudinal differences in the timing and duration of past fire intervals, including those occurring in recent centuries (Bianchi, 2000; Huber and Markgraf, 2003; Huber et al., 2004; Whitlock et al., 2007). Fire activity has been attributed to climate anomalies linked to the latitudinal position of the Southern Westerlies and the strength and location of the southeastern Pacific subtropical high-pressure system (Whitlock et al., 2007; Garreaud et al., 2008). In the mid-latitudinal region of Patagonia (latitude 38–438 S), periods of weakened and poleward-shifted Westerlies and a stronger than normal subtropical high-pressure system produce drought conditions and large areas burned as a result of higher than normal summer temperatures and lower than normal summer precipitation (Villalba, 1994; Kitzberger and Veblen, 2003; Whitlock et al., 2007). At interannual and interdecadal scales, seasonal moisture deficits and high fire occurrence are also related to El Niño–Southern Oscillation (ENSO) and decadal climate LATE QUATERNARY VEGETATION AND FIRE HISTORY variability. North of latitude 388 S, including the study site, winter and spring precipitation is somewhat enhanced during El Niño events (Compagnucci and Vargas, 1998; Montecinos and Aceituno, 2003), although it does not outweigh the moisture deficit resulting from higher than normal summer and autumn temperatures (Garreaud et al., 2008). Our research objective was to reconstruct the environmental history on the east side of the Andes at latitude 378 S at the northern limit of mixed Nothofagus obliqua/N. pumilio forest and the southern limit of the Monte desert scrub vegetation to better understand the climate history of this transitional region. We present pollen, charcoal and limnological data from Mallı́n Vaca Lauquen in northern Patagonia. Two time periods are of special interest: the Lateglacial and early Holocene interval (16 000–8000 cal. a BP) when summer insolation was lower than at present and ENSO variability was less pronounced (e.g. Markgraf and Diaz, 2000; Moy et al., 2002), and the late Holocene interval (last 3000 a) when ENSO variability was strengthened. Site description Mallı́n Vaca Lauquen is located near the confluence of the valleys of Lagunas Epulauquen and Vaca Lauquen about 60 km 249 north-west of the town of Andacollo, northwestern Neuquén Province (Fig. 1). The lakes are dammed by recessional moraines, the lowest of which lies at about 1500 m elevation. The surrounding peaks are about 2500 m elevation. The abundance of shrubs and introduced herbs in the local vegetation attests to heavy livestock grazing. The present-day vegetation consists of impoverished high-elevation bunchgrass steppe (estepa graminosa; León et al., 1998) with Festuca pallescens and shrubs in the families Asteraceae (Baccharis ssp., Chiliotrichium rosmarinifolium, Chuquiraga ssp., Mutisia ssp., Perezia ssp.) and Rhamnaceae (Colletia spinosissima), as well as Berberis rosmarinifolia, Ephedra frustillata and Eryngium paniculatum (Apiaceae). Native herbs include Osmorhiza berteroi (Apiaceae), Calceolaria biflora (Scrophulariaceae), Quinchamalium chilense (Santalaceae) and Phacelia sp. (Hydrophyllaceae), among others. Introduced herbs include Rumex acetosella and Plantago lanceolata. The mountain slopes are covered by the northernmost extent of southern beech (Nothofagus) forest, which is found up to treeline at 1700 m elevation. Nothofagus obliqua dominates lower elevation forests, while N. pumilio and N. antarctica grow at higher elevations, forming a krummholz belt above the treeline in the transition zone to the Andean tundra. N. antarctica also grows on poor soils at all elevations. Few Austrocedrus chilensis trees grow at lower elevations, and burnt and dead trees of Nothofagus obliqua in the watershed offer abundant evidence of fires in recent times. Figure 1 Location of the Mallı́n Vaca Lauquen site, showing major topographic features and vegetation cover. Nothofagus forests are in green; the steppe areas are in light brown, Andean tundra in grey Copyright ß 2008 John Wiley & Sons, Ltd. J. Quaternary Sci., Vol. 24(3) 248–258 (2009) DOI: 10.1002/jqs 250 JOURNAL OF QUATERNARY SCIENCE Given the absence of meteorological stations in this mountainous region, mean annual precipitation and temperature were estimated at 700 mm and 48C, respectively, based on data from the Climatologic Atlas of South America (J. A. Hoffman, ed.), UNESCO, 1975. Precipitation occurs primarily during autumm and winter months (April through July). A review of the climate of Patagonia and its relation to major vegetation zones lists mean annual precipitation at the steppe– forest ecotone of about 600 mm and mean annual temperature of 68C, based on 62 stations between latitudes 378 and 558 S (Paruelo et al., 1998). Methods In 2002, three sediment cores, MVL-02A, MVL-02B, and MVL02C, were taken with a modified Livingstone corer from the centre of the wetland. The most complete core, MLV-02C, was 342 cm long. In the laboratory, cores were split and the sediments described. The core MVL-02A was analysed for the upper 50 cm (this segment was not compacted compared to the same segment from core MVL-02C), and core MVL-02C was analysed between 50 and 342 cm depth. Magnetic susceptibility (MS; Sandgren and Snowball, 2001), was measured using a Bartington MS meter with a hand-held MS2E sensor. Measurements were taken every 0.5 cm along the entire length of core. For pollen analysis, 1 cm3 volumetric samples were taken at 2.5 or 5 cm intervals and prepared with standard techniques (Faegri and Iversen, 1989). Pollen grains were identified at 400 and 1000 magnification, with counts ranging from 150 to 400 grains. Terrestrial pollen percentages were based on the total sum of pollen grains from trees, shrubs and herbs and used to calculate the percentages of terrestrial and aquatic palynomorphs. Excluded from the terrestrial pollen sum were Cyperaceae, fern allies (mostly undifferentiated Polypodiaceae and a few Hymenophyllum and Pteris), aquatic taxa (Myriophyllum) and the algae Pediastrum boryanum var. longicorne and Botryococcus. Escallonia and Caryophyllaceae were also excluded from the pollen sum, given their erratic and probably local occurrences in high numbers in some intervals. Escallonia is a shrub that often grows at the margin of wetlands and Caryophyllaceae (locally present taxa are within the genera Stellaria and Arenaria pollen type) grow primarily on disturbed sandy–gravelly soils. Lycopodium tracer spores were added to each sample for calculation of pollen concentration (grains cm3). Changes in pollen percentage and concentration were used to interpret past vegetation changes supported also by CONISS cluster analysis (Grimm, 1987). Pollen data were plotted using TILIA programs (Grimm, 1992). A total of 72 pollen, spore and algal types were identified. Local tree taxa included Nothofagus dombeyi type (this pollen type includes N. dombeyi, N. betuloides, N. pumilio and N. antarctica; however, N. dombeyi and N. betuloides do not occur north of latitude 378 450 S). In this record, this pollen type probably reflects N. pumilio and N. antarctica. Other local tree pollen taxa are Nothofagus obliqua type (N. obliqua, N. procera) and Prumnopitys andina; rainforest taxa from the lowlands west of the Andean crest included Hydrangea, Myrtaceae and Weinmannia; and local steppe shrub taxa consisted of Lomatia/Gevuina type, Rhamnaceae, Berberis, Ribes and Verbena. Common steppe herbs were Acaena, Chenopodiaceae, Wendtia, Plantago and Phacelia. Wetland herbs (Ranunculaceae, Valeriana, Lamiaceae, Liliaceae, Gentiana) and introduced European taxa (Rumex, Plantago lanceolata) were also present. Macrofossil remains were Copyright ß 2008 John Wiley & Sons, Ltd. identified (Table 1) and total number of oospores of charophytes (Charales) plotted in the pollen diagram. Microscopic charcoal particles (larger than 20 mm) were counted on the pollen slides and converted to charcoal concentration (particles cm3). These data were converted charcoal accumulation rates (CHAR; particles cm2 a1) by dividing concentration by the deposition time of each sample (cm a1). In addition, samples of 1 cm3 volume were taken at 0.5 cm intervals for macroscopic charcoal and plant macrofossil analyses. These samples were washed through nested metal screens of mesh sizes of 250 and 125 mm. The residues were identified and counted under a stereo microscope. Because both macroscopic charcoal size fractions showed similar trends through time, charcoal counts were combined and divided by sample volume to calculate charcoal concentration (particles cm3). Charcoal concentrations were resampled into contiguous 25 a bins (the median resolution of the record) in order to sample over equally spaced time intervals through the record. CHAR was determined by dividing concentrations (particles cm3) by deposition time (a cm1) using Char Analysis software (Higuera et al., 2008). The slowly varying trend often referred to as background charcoal (BCHAR) was determined with a 700 a lowess smoother, robust to outliers. Throughout the paper, we distinguish between microscopic CHAR from discontinuous pollen slides and macroscopic CHAR from the high-resolution analysis of contiguous samples. Charcoal and terrestrial plant remains used for AMS radiocarbon dating were washed thoroughly with distilled water. Results Sediment description and chronology From the base at 342–220 cm depth, the sediment was composed of laminated inorganic silty clay that above 250 cm depth became more organic. Between 220 cm depth and the surface, the sediment consisted of compact peats with darker and lighter horizons. The magnetic susceptibility record (Fig. 3) indicates high magnetic susceptibility associated with inorganic sediment below 250 cm depth and low magnetic susceptibility above that depth in organic sediments. Peaks in magnetic susceptibility corresponded to volcanic ash layers, including three thick layers at 106.5–112 cm, 265.5–272.5 cm and 315.5–320.5 cm depth. Core depth was initially adjusted for minor sediment compaction (of 2–4 cm/50 cm core segment) by multiplying the compacted length by a correction factor (core length/ compacted length) to reconstruct the actual length of each core segment. For the purpose of developing an age model, core depth was further adjusted by excluding the three thick and pumiceous volcanic ash layers (106.5–112 cm, 265.5– 272.5 cm and 315.5–320.5 cm original depth, Fig. 3), assumed to represent rapid deposition. Hereafter, depth refers to adjusted depth. Eight AMS radiocarbon dates were obtained on sieved charcoal and organic material (Table 2 and Fig. 4). Two of the radiocarbon dates (AA-57612: 7790 65 14C a BP, 309–311 cm original depth; and AA-65184: 3751 39 14C a BP, 145– 145.5 cm original depth) were out of sequence and not used in the age model. Two other dates overlapped (AA-57613: 5162 14C a BP at 80.5 cm and AA-65183: 4505 14C a BP at 99 cm original depth), and probability calculations using BCal (Buck et al., 1999) indicated that the older date was an outlier. J. Quaternary Sci., Vol. 24(3) 248–258 (2009) DOI: 10.1002/jqs LATE QUATERNARY VEGETATION AND FIRE HISTORY 251 Table 1 List and counts of terrestrial and aquatic plant macrophytes and Cladocera remains identified (numbers cm3 volume) Depth (cm) Nitella opaca Nitella hyalina Characf. braunii Hypericum sp. Bryophyte spores Utricularia sp. Cladocera Zones 2 4 6 6 10 8 0 5 0 0 0 0 1 6 4 0 5 6 0 0 0 0 0 7 5 20 20.5 28.5 29.5 12 22 40 18 18 13 34 13 0 0 0 0 0 0 0 0 15 0 15 18 0 0 0 0 0 2 6 0 4c 34.5 47 58.5 59.5 60 10 18 10 10 23 9 0 8 4 44 0 0 0 0 0 0 0 0 0 0 15 15 10 10 12 0 0 0 0 0 0 0 0 0 0 76.5 80.5 81.5 91.5 95 96 96.5 12 21 2 22 3 15 22 17 36 31 46 34 59 31 0 1 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 15 0 0 0 0 0 0 0 0 0 0 1 2 8 5 0 1 4a 115 0 13 0 9 0 0 0 3 214 216 235 239 239.5 28 42 31 43 84 0 0 0 0 0 0 0 0 0 0 3 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 321 16 0 0 18 0 3 0 1 Hence, only five dates were used to develop the age model after conversion to calendar years before present (cal. a BP) using CALIB 5.0 (Stuiver et al., 2005). The dates were calibrated using Southern Hemisphere calibration to the date 4504 cal. a BP and Northern Hemisphere calibration for the dates of 8171, 10 250 and 13 820 cal. a BP. The basal age of the core was extrapolated to ca 16 000 cal. a BP. A range of possible calibrated dates and probability distributions were determined for every radiocarbon date 4b using CALIB 5.0.1 (Stuiver et al., 2005). Monte Carlo sampling was used to generate a cubic smoothing spline through all the dates 2000 times, and the final age–depth model was based on the median of all the runs (Higuera, 2008). Based on this chronology, deposition times were about 30– 40 a cm1 between ca. 17 000 and 7000 cal. a BP, 10– 20 a cm1 between 7000 and 4000 cal. a BP, 10 a cm1 between 4000 and 2200 cal. a BP, 30–40 a cm1 between 2200 and 400 a cm1 and 10 a cm1 during the last 400 a. Figure 2 Map showing sites of charcoal records (see Fig. 7) Copyright ß 2008 John Wiley & Sons, Ltd. J. Quaternary Sci., Vol. 24(3) 248–258 (2009) DOI: 10.1002/jqs 252 JOURNAL OF QUATERNARY SCIENCE Table 2 List of radiocarbon dates, corresponding calibrated ages and information on materials dated Lab code Depth (cm) 14 C age SD d13C% AA65182 AA57613 AA65183 AA65184 AA57611 AA65185 AA57612 AA65186 13 30.5–31 80.5–81.5 99–100 145–145.5 228.5–229 254.5–256 309–311 331–332 400 2740 5 162a 4 505 3 751a 8 171 10 250 7 790a 13 820 130 77 60 39 72 58 65 76 25 28.44 22 27.5 28.18 21.8 28.53 14.4 a cal age BP (median probability) 400 2 873 5 831 5 155 4 113 9 133 12 002 8 568 16 465 Upper Lower Material dated 2 5 5 4 9 12 8 16 2 5 4 3 8 11 8 16 European taxa Sedge fragments Sedge fragments/charcoal Sedge fragments/charcoal Sedge fragments Organic fragments Bulk sediment Organic fragments Bulk sediment 990 925 135 130 142 192 636 669 700 837 966 924 984 754 397 247 Dates not used in chronology. Pollen and charcoal record Throughout the record, the pollen assemblages are codominated by steppe and forest taxa, suggesting that, like at present, steppe grew in the valleys and Nothofagus forest grew on the mountain slopes. The pollen stratigraphy was divided into five zones using a constrained cluster analysis (Grimm, 1987) of terrestrial and aquatic taxa (Fig. 5). Microscopic CHAR provided a reconstruction of regional fire conditions, and macroscopic CHAR was used to infer local fire conditions (Fig. 6). The variable sedimentation rates suggest a changing depositional environment and, for that reason, charcoal peaks or particular fire episodes were not identified. Instead, total macroscopic CHAR was used to infer general fire activity during the time represented by different pollen zones. Figure 3 Magnetic susceptibility of the Mallı́n Vaca Lauquen sediment core (original depth) Copyright ß 2008 John Wiley & Sons, Ltd. Zone 1 (320–288 cm depth; 17 000–14 900 cal. a BP) featured 60–80% non-arboreal taxa, composed of Poaceae (40%); steppe shrubs, including Asteraceae (10–15%), Ephedra (2%), Rhamnaceae (5%) and Ericaceae (5–10%); and steppe herbs (15–20%). Arboreal taxa Nothofagus dombeyi type and Prumnopitys andina accounted for 10–20% each. N. obliqua type and rainforest taxa were present in trace amounts. Escallonia accounted for 5%. Pollen concentration was low. with a mean of 400 grains cm3. Pediastrum boryanum var. longicorne was abundant but fluctuated markedly (200– 1000%). At levels where Pediastrum boryanum var. longicorne was less abundant, traces of Myriophyllum, rare Utricularia seeds and oospores of charophytes were found. The charophyte was identified as Nitella opaca (Table 1). Microscopic and macroscopic CHAR values were negligible, suggesting that fires were virtually absent. The primary pollen constituents of this zone suggest herbaceous grassland with open ground. A considerable number of taxa were present that grow on sandy and rocky substrates, such as alluvial fans and cobble shorelines (Ephedra, Ericaceae (Pernettya), Plantago, Colobanthus (Caryophyllaceae)). These taxa are often found in Lateglacial basal samples, reflecting colonisation of disturbed terrain (e.g. M. Pollux, Markgraf et al., 2007) and thus the assemblage has no direct analogue with modern steppe assemblages (Paez et al., 2001; Markgraf et al., 2002). Despite the relatively high inorganic content the deposition time was slow (30 a cm1). Zone 2 (288–212.9 cm depth; 14 900–8600 cal. a BP) had slightly higher values of Poaceae than before (up to 50%). Nothofagus dombeyi type increased to 30%, whereas Prumnopitys andina decreased to 10%, and N. obliqua type continued to be present in traces, reaching 2–5% at some levels. Rainforest taxa (Hydrangea, Myrtaceae, etc.) were more prominent during this interval. Pollen abundance of steppe shrub taxa continued as before with a mean of 5%. Pollen of open ground taxa (e.g. Plantago) decreased to negligible amounts. Wetland herb types increased to 5%. Pollen concentration continued to be low with 300– 400 grains cm3. Pediastrum boryanum var. longicorne fluctuated around a mean of 200% and Nitella opaca appeared in the upper part of this zone (ca. 10 800 cal. a BP) with numbers between 20 and 50 cm3. At those levels, Pediastrum boryanum var. longicorne was rare or absent. Microscopic and macroscopic CHAR values were low in this zone but higher than before, implying some fires in the region. The deposition time was even slightly slower than in Zone 1 (40 a cm1). The assemblage indicates a more diverse herbaceous steppe than before, with less open and disturbed ground, comparable to the palynologically defined ’mid-grass steppe’ of Paez et al. (2001) J. Quaternary Sci., Vol. 24(3) 248–258 (2009) DOI: 10.1002/jqs LATE QUATERNARY VEGETATION AND FIRE HISTORY 253 0 Age (cal yr BP) 2000 4000 6000 8000 10000 12000 14000 cubic spline interpolant, 95% CI 16000 350 300 250 200 150 100 50 depth below mud-water interface (cm) 00.02 0.0810 30 50 sed. rate resolution (cm yr -1) (yr sample-1) Figure 4 Age–depth model and sedimentation rates for the Mallı́n Vaca Lauquen record or estepa graminosa (León et al., 1998). Reduced erosion, inferred from the decreased magnetic susceptibility of the sediments, could explain the markedly lower sedimentation rates than in the previous zone. Nothofagus forest along the mountain slopes, probably composed of both N. pumilio and N. antarctica, became denser, limiting the habitat for Prumnopitys andina. Zone 3 (212.9–107.5 cm depth; 8600–5300 cal. a BP) showed an increase of Nothofagus dombeyi type to 40%, on account of a decrease of Poaceae (40–50%). Prumnopitys andina decreased to trace amounts. Nothofagus obliqua type appeared continuously, reaching values of 5% in the upper part of the zone; steppe shrub pollen types continued with low values; and rainforest pollen percentages decreased. Wetland herb taxa slightly increased to 10% and Escallonia showed several peaks of 15%. Pollen concentration continued low with 400 grains cm3, except for one high value at 110 cm depth. High percentages of Cyperaceae (500–600%) indicate initiation of wetland conditions, although some open water must have existed intermittently to support the presence of Myriophyllum, oospores of Nitella opaca and Botryococcus in some levels. In the upper portion of this zone, Caryophyllaceae pollen (Stellaria/Arenaria pollen type) appeared with values between 100% and 500%, suggesting shoreline disturbance by seasonal water-level fluctuations. Both macroscopic CHAR and microscopic charcoal values increased in this zone. Macroscopic CHAR reached 0.80 particles cm2 a1, suggesting local fires, and microscopic CHAR was moderate (100 particles cm2 a1), Figure 5 Pollen record of Mallı́n Vaca Lauquen Copyright ß 2008 John Wiley & Sons, Ltd. J. Quaternary Sci., Vol. 24(3) 248–258 (2009) DOI: 10.1002/jqs 254 JOURNAL OF QUATERNARY SCIENCE Figure 6 Charcoal record of Mallı́n Vaca Lauquen consistent with more regional fire activity. Deposition time was twice as fast (10–20 a cm1) as during zones 1 and 2, reflecting the onset of peat growth. Zone 4 (107.5–13 cm depth; 5300–400 cal. a BP) is characterised by the onset of continuing presence of Nothofagus obliqua type together with high values of N. dombeyi type. Rainforest and steppe shrub pollen types were sporadically present. Pollen of steppe herbs and wetland herbs were less abundant overall compared with values in Zone 3. This zone can be divided into three subzones based on changes in the proportions of arboreal and non-arboreal taxa. The initial subzone (4a: 107.5–60.2 cm depth; 5300–4200 cal. a BP) had 50% Poaceae, 30% Nothofagus dombeyi type, as well as slightly higher percentages of Acaena and lower Asteraceae. Pollen concentration continued low with 400 grains cm3, although deposition time was high, as in Zone 3 (10– 20 a cm1). The middle subzone (4b: 60.2–30.8 cm depth; 4200–2200 cal. a BP) showed slightly lower values for Poaceae (40% mean) and steppe taxa and slightly higher values for Nothofagus dombeyi type (45%). Pollen concentration doubled with values of 800 grains cm3, reaching a maximum of 1200 grains cm3, while deposition time continues high (10 a cm1). The upper subzone (4c: 30.8–13 cm depth; 2200– 400 cal. a BP) featured an increase in Poaceae percentages to 50% and a decrease of Nothofagus dombeyi type to 40%. Pollen concentration was slightly lower with 700 grains cm3 and deposition time was slower as well (30–40 a cm1). Cyperaceae decreased to a mean of 200%, whereas aquatic taxa increased. Among the aquatic taxa, Charophyte oospores were persistently high, although highly variable, especially in subzones 4a and 4c and one level in 4b. Charophytes were dominated by Nitella hyalina (Table 1), a shallow-water species that currently grows in 0.15–0.8 m water depth (Garcı́a, 1987; Cáceres and Garcı́a, 1989); N. opaca, a species tolerating deeper waters, was also present in low amounts. At one level in Copyright ß 2008 John Wiley & Sons, Ltd. subzone 4a, Chara braunii, a taxon commonly associated with Nitella hyalina (Corillion, 1957), was found. Pediastrum boryanum var. longicorne appeared intermittently with low values in subzones 4a and 4c; Myriophyllum was essentially absent except for occurrences in subzone 4a. Botryococcus had rare occurrences. Subzones 4a and 4c indicate shallow, fluctuating water levels; subzone 4b had high amounts of bryophyte spores, indicating intermittently dry conditions. During 4a, macroscopic and microscopic CHAR reached their highest levels of the record (macroscopic CHAR was up to 12.0 particles cm2 a1 and microscopic CHAR reached 1516 particles cm2 a1). These high levels decreased dramatically in subzone 4b, and after 3500 cal. a BP very few fires were registered. In subzone 4c, CHAR levels were again elevated (macroscopic CHAR was up to 14.50 particles cm2 a1 and microscopic CHAR reached 6650 particles cm2 a1), indicating significant local and regional fire activity. The pollen and charcoal data suggest that moisture variability must have been higher overall, especially in the lower and upper subzones (4a and 4c) than in the middle subzone to create conditions for Nothofagus obliqua type to occur, and higher fire activity. Overall conditions, however, were wetter than in Zone 3, especially in subzones 4a and 4c. Zone 5 (13–0 cm depth; 400–0 cal. a BP) showed a marked decrease in Poaceae, coupled with an increase in Nothofagus dombeyi type, steppe shrub taxa and mesic herbs. Introduced European weed taxa appeared (Plantago lanceolata, Rumex acetosella), indicative of grazing and logging. During the upper part of this zone, Nothofagus dombeyi type also decreased, documenting more intense forest clearance in more recent times. Pollen concentration continued high with 800 grains cm3 and deposition time increased (10 a cm1), suggesting higher erosional input. The charophytes, Nitella hyalina and N. opaca, dominated during this interval, while Pediastrum boryanum var. longicorne and Myriophyllum were present intermittently, indicating seasonally fluctuating water levels, as observed during fieldwork in 1981. CHAR values in the last 1000 a are relatively low (macroscopic CHAR was <0.90 particles cm2 a1 and microscopic CHAR was <530 particles cm2 a1). These data suggest a limited role for fire in the local watershed as well as regionally in recent centuries. Palaeoenvironmental and palaeoclimate history The general vegetation history inferred from this core is comparable to the early study (Markgraf, 1987). However, the additional detailed record of limnological changes and fire history provide a more detailed palaeoenvironmental and palaeoclimatic picture. The different proportions of aquatic taxa in the two records are related to the fact that the present record was taken in the deepest, wettest part of the basin, which was dry in 2002 but inundated in 1981. The earlier core, taken at a marginal location, showed high percentages of Botryococcus and low values of Pediastrum boryanum var. longicorne, indicating shallow and fluctuating water levels. The present record has high amounts of Pediastrum boryanum var. longicorne and only a few levels with Botryococcus, reflecting its deeper water location. During the last 17 000 cal. a, open-water conditions must have existed in the shallow basin, allowing for the continuous presence of aquatic taxa. Between 16 000 and 14 800 cal. a BP, high amounts of aquatic taxa were present: Pediastrum J. Quaternary Sci., Vol. 24(3) 248–258 (2009) DOI: 10.1002/jqs LATE QUATERNARY VEGETATION AND FIRE HISTORY boryanum var. longicorne, an algae common in small, highelevation lakes (Komárek and Fott, 1983; Jankovská and Komárek, 2000); the charophyte Nitella opaca, a cosmopolitan species that thrives in water depths of 0.5–2.5 m, but also grows in clear waters up to 40 m depth (Corillion, 1957); Utricularia, a montane to subalpine, shallow-water plant; and shallow-water diatoms growing in low-productivity environments (Fragilaria construens, var. venter and Cyclotella stelligera in the basal samples of the original core; Markgraf, 1987). These taxa suggest clear, dystrophic and slightly acidic waters. Such lowproductivity environments were apparently common in the Lateglacial period during the early stages of lake development (Bradbury and Whiteside, 1980). The initial terrestrial vegetation at Mallı́n Vaca Lauquen between 17 000 and 14 900 cal. a BP was a shrub–steppe with considerable open, sandy and gravely ground. This patchy landscape suggests cool and dry conditions. An interesting aspect of this interval is the occurrence of Prumnopitys andina pollen, which is also documented at the Tagua Tagua site west of the Andes at latitude 34.58 S (Heusser, 1990). Prumnopitys does not occur at either site today, but scattered individuals grow on the Chilean side of the Andes from latitude 35.58 S to 438 S at 600 and 1000 m elevation, in association with Austrocedrus chilensis, Nothofagus obliqua, N. pumilio and Lomatia hirsuta (Veblen et al., 1995). At latitude 388 S to the south of Mallı́n Vaca Lauquen, Prumnopitys andina crosses the Andes into Argentina as individual trees near upper treeline (Donoso, 1974). This limited geographic and elevational distribution is confined to areas of low temperatures and dry summers (Veblen et al., 1995). Apparently, Prumnopitys andina was more widespread than today on both sides of the Andes during the initial period of deglaciation (Markgraf et al., 1992, 2002; Whitlock et al., 2001). Its presence at Mallı́n Vaca Lauquen is consistent with about 68C cooler Pacific sea surface temperatures (latitude 418 S, Lamy et al., 2004; latitude 348S, Kim et al., 2002), a strengthened southeastern Pacific highpressure system, and weakened Southern Westerlies (Markgraf et al., 1992; Lamy et al., 1999). Between 14 900 and 8600 cal. a BP, the steppe environment was more diverse than before and the high proportion of herb pollen is characteristic of modern pollen assemblages from the present-day grass steppe (Paez et al., 2001), where precipitation is about 500 mm (Paruelo et al., 1998). Forests on the slopes around the site continued open with Prumnopitys andina and Nothofagus pumilio/N. antarctica. Higher amounts of longdistance pollen from rainforest taxa are attributed to an expansion of Valdivian rainforest in the Chilean lowlands (Moreno, 1997, 2004). Initially the lake was still dystrophic but increased abundance of the charophyte Nitella opaca by 10 000 cal. a BP suggests increased alkalinity. Conditions were warmer and wetter than before, although the openness of the forest at Mallı́n Vaca Lauquen suggests that precipitation was less than at present. Fires were not a significant component of the ecosystem, judging from low CHAR values. Drier-than-present conditions have also been suggested from marine and terrestrial records at latitude 348 S (Lamy et al., 1999; Villa-Martı́nez et al., 2004). In contrast, records between latitudes 40 and 458 S show rainforest expansion by 14 800 cal. a BP, suggesting that moisture had reached present levels in that region (e.g. Moreno, 1997, 2004; Villagrán, 2001; Markgraf et al., 2002; Haberle and Bennett, 2004). Westerly winter storms had apparently become established south of latitude 408 S at this time (Markgraf et al., 2002; Whitlock et al., 2006), but the southeastern Pacific High continued to be stronger than today. This combination resulted in a less pronounced precipitation increase to the north of latitude 408 S than to Copyright ß 2008 John Wiley & Sons, Ltd. 255 the south (Markgraf et al., 1992; Lamy et al., 1999; Romero et al., 2006). After 12 000 cal. a BP, however, aridity returned to mid latitudes. This early Holocene (12 000–8000 cal. a BP) aridity has been widely documented in marine and terrestrial records between latitudes 54 and 338 S (Villagrán, 1991, 2001; Markgraf et al., 1992, 2007; Lamy et al., 1999, 2001, 2004; Huber et al., 2004; Moreno, 2004). In the rainforests west of the Andes, drought- and disturbance-adapted rainforest elements (especially Nothofagus obliqua and Weinmannia trichosperma) became dominant at that time (Villagrán, 1991, 2001; Moreno and León, 2003; Abarzúa and Moreno, 2008) and high-latitude Nothofagus forests continued quite open (Huber et al., 2004). The duration of this interval is longer and more pronounced at high latitudes than at mid latitudes and east of the Andes, as illustrated by a comparison of charcoal records between latitudes 52 and 358 S (Fig. 7). Highest fire activity at latitude 528 S occurred between 12 000 and 5500 cal. a BP and at 458 S between 11 000 and 7000 cal. a BP (Rio Rubens: Huber et al., 2004; Mallin Pollux: Markgraf et al., 2007). At latitude 42–358 S, west of the Andes, highest fire activity occurred between 11 000 and 9000 cal. a BP and after 3000 cal. a BP (Lago Melli: Abarzúa and Moreno, 2008; Tagua Tagua: Heusser, 1990; Purén-Lumaco: Abarzúa, pers. comm.). East of the Andes at lat 418 S (Laguna el Trébol: Whitlock et al., 2006), high activity is registered after 8000 cal. a BP with 1000 a long maxima centred at 6500, 4500, 3500 and 1000 cal. a BP. Arid conditions during the early Holocene have been attributed to substantially weaker Westerlies or latitudinally shifted westerly storm tracks, related to a strengthened southeastern Pacific High (Markgraf et al., 1992; Whitlock et al., 2007). However, a poleward shift in the Westerlies does not fully explain these east–west differences in fire activity. An anomalous component of upper-level flow associated with the west–east dipole may have resulted in opposite moisture regimes west and east of the Andes, but this hypothesis bears further study. Mallı́n Vaca Lauquen did not show this early Holocene arid and warm interval with high fire activity; instead conditions continued cool and dry until 8600 cal. a BP. Only between 8600 and 5300 cal. a BP did conditions became markedly warmer at Mallı́n Vaca Lauquen, based on the development of Cyperaceae wetland in the shallow basin, only briefly interrupted by periods with open water. Precipitation became more variable, as evidenced by fluctuations in local shoreline taxa (Escallonia, Arenaria/Stellaria pollen type) and between steppe and forest taxa. The steppe assemblage compares with present-day shrub steppe (estepa arbustivo-graminosa; León et al., 1998, Paez et al., 2001), where precipitation is <400 mm (Paruelo et al., 1998). Elevated levels of microscopic CHAR at Mallı́n Vaca Lauquen, especially between 5000 and 4500 cal. a BP, may have come from regional fires in the Chilean lowlands. The Tagua Tagua site features high levels of charcoal at this time (Heusser, 1990) (Fig. 7). Higher levels of charcoal and low lake levels are also observed between ca 10 000 and 6000 cal. a BP in a record from the Purén-Lumaco valley (latitude 388 300 S, longitude 738 W), located on the southeastern slope of the Coastal Chilean Range (Nahuelbuta Range, Abarzúa, pers. comm.). The macroscopic charcoal record suggests some local fires at Mallı́n Vaca Lauquen at about 7500 cal. a BP. By 5300 cal. a BP, the present-day mixed Nothofagus obliqua/N. pumilio forest had become established in the region, indicating a shift to present-day winter rain/summer drought conditions. High abundance of charophytes, Nitella opaca and N. hyalina at Mallı́n Vaca Lauquen suggests more permanent water than before. Nitella hyalina is a shallow-water species growing at 0.15–0.8 m water depth (Garcı́a, 1987; J. Quaternary Sci., Vol. 24(3) 248–258 (2009) DOI: 10.1002/jqs JOURNAL OF QUATERNARY SCIENCE Ri o R la ub t 5 en 2. s 13 S, lo ng 71 .8 8W M ,e al lin le v. la P 22 t 4 ol 0 5. lux m 08 S, lo ng 71 .8 La 4W go ,e la M le t 4 el v. l 64 2. i 76 0 m S, lo ng 73 .5 5W ,e Ta le gu v. 70 a la Ta m t3 g u 5. a 50 S La gu , lon g la n a 71 t 4 el .1 5. Tr 6W 08 eb ,e S, ol le lo v. ng 20 71 0 .8 m 4W ,e M le al v. la lin V 75 t3 a 8 6. ca m 86 L a S, u lo qu ng e 71 n .0 5W ,e le v. 15 67 m 256 Age (cal yr BP) 16.2 0 0 50 100 150 Particles cm-2 yr-1 Huber et al. 2003 0 4 Particles cm-2 yr-1 8 0 Markgraf et al. 2007 10 20 Particles cm-2 yr-1 Abarzua & Moreno 2008 Particles x 103 gm-1 Particles cm-2 yr-1 0 Particles cm-2 yr-1 Heusser 1990 Whitlock et al. 2006 this study Figure 7 Charcoal records for sites in southern South America (for location of sites see Fig. 2) Cáceres and Garcı́a, 1989), whereas N. opaca tolerates deeper waters, generally around 2 m, but occasionally to 40 m depth (Corrillion, 1957). The fluctuations between these two taxa indicate variable conditions, with shallower water between 5300 and 4200 cal. a BP and after 2200 cal. a BP and slightly deeper water between 4200 and 2200 cal. a BP. The limnological changes seen at Mallı́n Vaca Lauquen are more or less synchronous with fluctuations in the abundance of the wetland taxon Typha at the Tagua Tagua site during the last 5000 cal. a (Heusser, 1990). Freshwater algae at Laguna de Culeo during the last 2500 cal. a (Chilean lowlands at 348 S; Jenny et al., 2002; Villa-Martı́nez et al., 2004) are also interpreted to reflect increased moisture coupled with high hydrologic variability. Very high CHAR values between 5300 and 4200 cal. a BP and to a lesser degree after 2200 cal. a BP indicate increased fire activity locally and regionally, as a result of more fuel biomass and more suitable fire climate. Reduction of Nothofagus pumilio/N. antarctica forest cover and the expansion of grassland and disturbance taxa are consistent with more fire activity at this time. High charcoal abundance during this interval is also seen in records from the Chilean lowlands at Tagua Tagua (Heusser, 1990), the Purén-Lumaco valley (Abarzúa, pers. comm.) and Lago Melli (Abarzúa and Moreno, 2008), beginning by 3000 cal. a BP. Higher-than-present moisture coupled with increased multicentennial variability by 5000 cal. a BP has been inferred from marine and terrestrial records (McGlone et al., 1992; Lamy et al., 1999; Jenny et al., 2002; Villa-Martinéz et al., 2004). At latitudes north of 408 S, it is likely that ENSO variability and associated increased convective storm activity during El Niño years increased fire activity in the last 5500 cal. a (Kitzberger and Veblen, 2003) (e.g. Markgraf and Diaz, 2000; Moy et al., 2002). Multi-centennial records, such as those presented here, cannot document ENSO’s inter-annual and inter-decadal Copyright ß 2008 John Wiley & Sons, Ltd. variability. However, the present-day correlation between El Niño-related drought conditions (Montecinos and Aceituno, 2003) and historical fire years (Kitzberger and Veblen, 2003) suggests that ENSO variability is a factor in mid-latitude fire occurrence. Palaeoenvironmental records have indicated that ENSO variability was strengthened during the mid and late Holocene (McGlone et al., 1992; Moy et al., 2002), and strong ENSO variations may account for high fire activity at 5000 cal. a BP (primarily east of the Andes), 3000 cal. a BP and between 2000 and 1000 cal. a BP. One could speculate that high fire activity in the late Holocene seen similarly in the mid latitudes on both sides of the Andes reflects the influence of ENSO and the build-up of fuels. The last 400 cal. a show the influence of humans on the vegetation and fire regimes of the Mallı́n Vaca Lauquen region. A decrease in Poaceae and Nothofagus and an expansion of European weeds (Rumex acetosella, Plantago lanceolata) are noted in the pollen record and attributed to a decrease in grassland from livestock grazing and a decrease of Nothofagus due to forest clearance. The impact began in the 16th century with the arrival of Spanish colonists. The markedly faster deposition time after 400 a (30 a cm1) is probably not the result of greater in-wash of clastic sediments into the basin because magnetic susceptibility values are low. Instead, it may be the result of greater water run-off leading to more persistent moisture in the basin and wetland development. Low levels of charcoal at this time, implying negligible or small-scale burning, support the local wet site conditions. Conclusion High relief and steep climatic gradients in the southern Andes produce diverse environments that are also reflected by a J. Quaternary Sci., Vol. 24(3) 248–258 (2009) DOI: 10.1002/jqs LATE QUATERNARY VEGETATION AND FIRE HISTORY diverse environmental history. Mallı́n Vaca Lauquen represents the only high-elevation site so far studied with mixed Nothofagus obliqua/N. pumilio forest located near the transition to the Monte desert scrub vegetation. One unique feature of the past history is the Lateglacial and early Holocene (17 000–8600 cal. a BP) presence of Prumnopitys andina, indicating cooler and drier summers than today. This might perhaps explain the absence of fires in that area at this time. The late appearance of Nothofagus obliqua after 5000 cal. a BP represents another environmental change not seen in other records. Together with more permanent water in the lake, the presence of N. obliqua marks the establishment of the presentday winter rain/summer drought climate regime in the late Holocene. Although Austrocedrus chilensis is locally present in the forests in small numbers, it never played a major role in this region’s past history, in contrast to records farther south and east of the Andes. Unlike most sites in southern Patagonia, which show high fire activity in the early Holocene, Mallı́n Vaca Lauquen and Laguna el Trébol at latitude 41.58 S record high charcoal levels in the mid and late Holocene. At Mallı́n Vaca Lauquen, inferred fire activity is highest between 5300 and 4200 cal. a BP and after 2200 cal. a BP. The latitudinal differences in timing may be related to fundamentally different climatic controls on fire activity, in particular the stronger influence of ENSO variability at mid latitudes. Additional records are needed in this region to describe the geographic extent of this pattern and better understand the underlying drivers of past fire activity. Acknowledgements NSF grants to CW (ATM 0117160 and ATM 0714061) are acknowledged. 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