Abstract
Introduction
The mid- to late Holocene climate of the Southern Hemisphere high latitudes is characterised by an early Holocene warm period from 11,000 to 9500 cal. yr BP, a general cooling until 4500 cal. yr BP and a mid–late Holocene warm period from 4500 to 2800 cal. yr BP or later (Bentley et al., 2009). Following this, a cooler period generally persisted until the present rapid regional warming of the last few decades with many regional variations (Hodgson et al., 2009).
The mid–late Holocene warm period in the Antarctic Peninsula region has been attributed to increases in solar insolation (Bentley et al., 2009) and changes in the meridional position of the core of the Southern Hemisphere Westerly Wind (SHW) belt. Understanding the link between climate changes and the position and strength in the SHW in this region is important because changes in the winds can affect upwelling in the Southern Ocean, which in turn affects global CO2 levels (e.g. Anderson et al., 2009; Björck et al., 2012; Hodgson and Sime, 2010; Le Quéré et al., 2010; Watson and Garabato, 2006). Multiple climate model simulations (Varma et al., 2012) suggest an overall strengthening and poleward shifting trend in the SHW during the course of the mid- to late Holocene corresponding with a cooling trend in Antarctica (Masson et al., 2000). However, validating the model results with reconstructions of the past position and intensity of the SHW still remains elusive. For example, Varma et al. (2012) note that proxy records from western Patagonia (Moreno et al., 2010) suggest a trend of increasing SHW strength during the past 7000 years, which is not supported by sedimentological and pollen-based reconstructions of precipitation variability in South Patagonia (see Fletcher and Moreno, 2012; Lamy et al., 2010).
These studies highlight a limitation of many high-latitude palaeoclimate records which show considerable variability not only in the timing of inferred warm and cool periods (which is further compounded by low resolution age control in many studies; Table 1), but also in differences in the precise climate forcing mechanisms used to explain them. In the core belt of the SHW, the lack of reliable reconstructions is particularly acute because of the limited amount of landmasses from which palaeoclimate records can be obtained. These landmasses consist of the southernmost tip of South America and a small number of islands in the sub-Antarctic region. Even at these locations, local orographic, glaciological and edaphic factors can overprint the regional climate signals.
Summary of previously interpreted Holocene cooling and warming periods in the South Atlantic sub-Antarctic islands, maritime Antarctic Islands and Northern Antarctic Peninsula.
LOI: loss-on-ignition.
The number of published radiocarbon dates is shown for each record to indicate the density of age control. Non-calibrated radiocarbon dates have been calibrated using the calibration curve SHCal13 in CALIB 7.0.2 (Stuiver et al., 2014).
Here, we present a well-dated, high-resolution pollen record of mid- to late Holocene vegetation and climate change from a lake sediment core from Fan Lake, Annenkov Island, South Georgia (Figure 1). The South Georgia archipelago lies in the core belt of the SHW, and its climate is governed by the latitudinal movement of the Polar Front Zone (PFZ) and the SHW (Bentley et al., 2007), which makes it particularly sensitive to even relatively minor climatic fluctuations. Most existing palaeorecords have been taken from the north and northeastern side of South Georgia (Bentley et al., 2007; Rosqvist and Schuber, 2003), but sites off the southwest coast, such as Annenkov Island, are thought to be particularly responsive because they are not affected by active glaciation, or orographic influences prevalent on the northeastern sites.

(a) Location map of the South Atlantic, showing the location of the main frontal systems of the Southern Ocean influencing the climate of South Georgia, South America and the Antarctic Peninsula. Landmasses are shaded black; areas less than 500 m deep are shaded grey; ACC: Antarctic Circumpolar Current. (b) LANDSAT image of South Georgia, showing the location of Annenkov Island and place names mentioned in text. (c and d) Fan Lake field site. (e) Satellite image of Annenkov Island. Red colour indicates vegetation cover on 25 October 2012 calculated from very high-resolution Quickbird2 satellite images with an on-the-ground resolution of 2.4 m per pixel. Live green vegetation in each pixel of the satellite image shows up as red in the image because the cell structure of chlorophyll containing cells absorbs red light while reflecting light in the infra-red part of the spectrum (Fretwell et al., 2011).
The overall aims of our project were (1) to establish when Annenkov Island was last glaciated, (2) to assess whether the timing and extent of any periods of Holocene glacier advance were synchronous or offset with known phases of glacier advance on mainland South Georgia and (3) to test the hypothesis that Holocene palaeoenvironmental change on Annenkov Island is more directly coupled to changes in the SHW circulation compared with sites elsewhere on South Georgia that are influenced by orographic effects. The first two aims are addressed further in forthcoming papers. In this paper, our aims were to identify long-term regional changes in climate and vegetation and to test whether long-distance pollen influx at Fan Lake is related to previously reconstructed changes in SHW circulation.
Site and present day vegetation
South Georgia is located in the South Atlantic Ocean approximately 1300 km east–southeast of the Falklands Islands (Figure 1). The island is ~170 km long with mountains (Allardyce Range) reaching up to 2000 m down the middle of the island (Bentley et al., 2007; Rosqvist and Schuber, 2003). Meteorological records from King Edward Point show well-defined summer and winter seasons (Barrow, 1978) on the north coast of South Georgia with a mean annual temperature of 2°C (Van der Putten et al., 2004) and a mean annual precipitation of around 1400 mm at sea level (Rosqvist and Schuber, 2003). South Georgia is positioned at the maximum northern limit of Antarctic pack ice, and over half the island is permanently covered in glaciers and ice fields (Bentley et al., 2007).
Annenkov Island (54°29′S–55°00′S, 35°30′W–38°30′W; Figure 1) is situated
Cores were extracted from the deepest point of Fan Lake (18 m) and from the centre of a peat land area (henceforth referred to as KH4; 54°29.927′S, 37°02.876′W)
The vegetation in the region is dominated by sub-Antarctic herbs and shrubs such as
One potential problem of reconstructing past vegetation on South Georgia is the selective preservation of pollen produced by some species. For example, while the flowering plants,
South Georgia experienced little or no human activity until the establishment of sealing and whaling stations on the north coast of the island during the last
Material and methods
Sedimentology and chronology
Sediment cores were extracted from Fan Lake (Figure 1d) using UWITEC gravity and percussion driven piston corers. Two 18-cm UWITEC gravity cores were extracted to recover the near-surface sediments from the main core site and sectioned at 0.5-cm intervals, with a thick moss layer preventing deeper coring. The UWITEC piston corer was deployed to extract deeper sediments in a series of 2-m long offset drives capturing 1.87–1.90 m of sediment with 20 cm overlaps to produce a continuous sequence down to 5.8 m, where drilling was stopped by a large rock or bedrock. The cores were photographed (visual and x-radiographic imagery), macroscopically described and analysed for wet density, dry mass (105°C for 12 h), organic content (loss-on-ignition (LOI) at 550°C for 2 h) and carbonate composition (LOI at 925°C for 2 h, multiplied by 1.36 to provide a first approximation of total carbonate) following standard methods (Dean, 1974). Known overlaps, the main changes in lithology defined initially by density, LOI, Barington Instruments MS2G loop sensor (1 cm sensor, 10-s measurement time) at 2-mm intervals with standard calibration procedures (Gunn and Best, 1998), were used with the position of visible moss layers and key lithological markers, to stack the five core sections into a single stratigraphic sequence and to produce a composite depth sequence.
Lead-210 (210Pb) and Ceasium-137 (137Cs) dating analysis was undertaken on the upper 11 cm of the surface core using dried and homogenised samples packed into a 40-mm tube to 40 mm depth and left to stand for at least 21 days to allow 226Ra and 214Pb to reach equilibrium. Samples were measured on an Ortec J-shape ultra-low background germanium well detector and remote preamplifier. Data analysis and dating model calculations were undertaken following standard procedures as defined in Appleby (2001). 137Cs was at or below detection limits. Unsupported 210Pb estimates were derived from both the constant initial concentration (CIC) and constant rate of supply (CRS) method (Table S1, available online) (Appleby and Oldfield, 1978). CRS ages were used in whole core age–depth modelling.
A total of 32 samples for radiocarbon dating were taken from the Fan Lake core. One basal sample from the KH4 peat core was also dated to provide an additional constraint on the timing of catchment deglaciation (Table 2). Calibration of terrestrial radiocarbon ages was carried out in OXCAL v.4.2 (Bronk Ramsey, 2009) using the SHCal13 Southern Hemisphere atmosphere dataset (Hogg et al., 2013). For post-bomb samples, absolute percentage of modern carbon (pMC) data were corrected according to 13C/12C isotopic ratios from measured pMC, where a ‘modern’ pMC value is defined as 100% (
Radiocarbon age summary data for Fan Lake sediment core and the basal age from KH4 peat bog core.
pMC: percentage of modern carbon; M and TM: moss macrofossil and species name; BOM: bulk organic matter; BS: bulk sediment.
FANSC = UWITEC gravity surface core sliced into 0.5-cm subsamples in the field; FAN2A-1 is the first UWITEC core drive. The sediment–water interface was recovered for both FANSC and FAN2A-1, and excess surface water was pipetted off from the top of the FANSC core before subsampling in the field. The FAN2A-1 core was packed to minimise disturbance of the flocculent upper sediments during transit. Cores 2A, 2B, 1C, 1E were used to construct the composite downcore sequence shown in Figure 2.
All radiocarbon age data were included in age–depth modelling to test agreement with the following manually assigned rejection criteria: (1) age reversals where independent 210Pb dating evidence suggests radiocarbon ages are ‘too old’ or reversed, (2) ages from near tops of cores which could have been water washed or disturbed during coring, (3) moss fragments or strands which are more likely to be reworked and (4) improbable age reversals where macrofossil ages occur higher stratigraphically than younger macrofossils and are beyond two-sigma calibrated age range of lowermost macrofossil, or represent bulk inorganic sediment that is more than likely reworked.
Most likely age by comparison with Pb-210 data.
A master age–depth model was constructed using a combination of all 210Pb CRS ages and 32 radiocarbon ages in a Bayesian age–depth model undertaken in BACON v2.2 (Blaauw and Christen, 2011; Figure 2; Table 2). All ages quoted in text are weighted mean ages produced from the BACON age–depth model rounded to the nearest 10 years (Figure 2; Table 2).

(a) Age–depth model for calculation of down-core radiocarbon ages (see text) compared with visual and x-radiographic core imagery (taken using an ITRAX XRF core scanner; running conditions 50 kV, 50 mA, 200 ms, 100 µm measurement interval), sedimentology and grain size summary data for the composite Fan Lake core. X-radiographic images are negatives; hence, areas with greater relative density are grey–white and darker areas are more organic or, in Units 1-2, represent areas of the core where clasts were removed before scanning. Age–depth modelling was undertaken in BACON v2.2 (Blaauw and Christen, 2011) using all radiocarbon ages and constant rate of supply (CRS) age model 210Pb data (shown as white circles in c; see also Table S1, available online) using the SHCal13 calibration curve (Hogg et al., 2013) with 5-cm segments, 121 segments in total, for 27 million iterations and with every 615th iteration selected. Initial parameters were mean accumulation rate of 20 yr cm−1, accumulation shape = 1.5, mem. strength = 20. When tested, mem. strength values of 10–100 gave essentially the same result, increasing confidence in the model parameters chosen and the reliability of the model. Calibrated age probability distributions with identification numbers are shown in blue. The median best-fit age plot (1-mm interval) in red is superimposed on individual BACON data runs shown as a grey shaded area. Darker grey areas indicate a greater density of individual age–depth model runs. The solid grey line represents the 95% confidence age limits for model runs. Colour versions of the core image can be viewed in the online version of this article; (b) Set up parameters for the BACON age–depth model; (c) Pb-210 ages (±1-sigma errors) from the CRS model which were used in the whole core age–depth model (red line and grey shading as described in a; (d) Sedimentation rates for the Fan Lake core calculated at 1 cm and 2 mm intervals.
Palynological analysis
Below 92 cm, the core was subsampled at 8-cm increments for pollen analysis. To capture higher resolution changes during the late Holocene, the uppermost ~92cm of the core was subsampled at 3- to 4-cm increments.
Approximately 1–2 g of sediment per sample were run through a 125-µm sieve after treatment with KOH and HCl. Standard laboratory techniques (Fægri and Iversen, 1989) including HF treatment and acetolysis were used to process the pollen samples. A
The Tilia program was used to produce percentage and influx pollen diagrams (Grimm, 1990) with pollen assemblage zones delimited by stratigraphically constrained cluster analysis in CONISS (Grimm, 1987). The significance of the zones was assessed using a broken-stick model in the Rioja package (Juggins, 2012). The pollen sum that is used to calculate percentage pollen diagrams is the total of pollen, excluding fungal spores and pollen grains derived from long-distance transport.
Multivariate analysis of the pollen data
Principal components analyses (PCAs) of the Fan Lake pollen data were carried out in PRIMER 6 and PAST (Clark and Gorley, 2006; Hammer et al., 2001). Rare (consistently <1%) pollen taxa were excluded, and remaining pollen percentages were normalised in PRIMER 6 (Clark and Gorley, 2006) prior to analysis. Confidence intervals (95%) were calculated for the total long-distance taxa (
Results
Sedimentology and chronology
The 587-cm Fan Lake core lithology was divided into eight units (Figure 2), from a basal unit (Unit 1) composed of a clast-supported sandy matrix to the uppermost units, which consist of laminated organic lake muds (Unit 7), capped by a living microbial mat and lake mud deposit (Unit 8; Figure 2). Between
Reworking of older material appears to have affected some of the bulk ages from glacially derived sediments near the base of the core. Within Unit 7, reworking of ‘older’ moss layers is evident between
Palynology
Pollen grains are well preserved through the core with influx continually high between approximately 350 and 150 cm. The major taxa found throughout all the pollen zones are
Three major pollen zones (Fan1, Fan2 and Fan3a–e) were defined by the stratigraphically constrained cluster analysis. Fan1, Fan2 and Fan3d–e were statistically significant as assessed using the broken-stick analysis. Other zones defined by the stratigraphically constrained cluster analysis were not significant but retained to aid description and interpretation of the data (Figure 3).

Pollen percentage diagram with pollen zones based on CONISS subdivisions (Grimm, 1987). Taxa with low percentages have a 10-times exaggeration from the second line in order to better show changes in abundance, and therefore
Pollen zone Fan1: 587–340 cm; ~7700–3790 cal. yr BP
Poaceae and

Summarised pollen influx diagram (grains cm−2 yr−1) showing most relevant pollen taxa.
Pollen zone Fan2: 340–232 cm; ~3790–3050 cal. yr BP
A drop in Poaceae to 65–80% is recorded while
Pollen zone Fan3: 232–0 cm; ~3050 cal. yr BP to present
The pollen zone is characterised by lower
Pollen sub-zone Fan3a: 232–200 cm; ~3050–2750 cal. yr BP
Increasing
Pollen sub-zone Fan3b: 200–154 cm; ~2750–2430 cal. yr BP
Poaceae percentages remain from 70% to 80% through this zone, and
Pollen sub-zone Fan3c: 154–94 cm; ~2430–1670 cal. yr BP
Poaceae percentages are on average higher compared with zone Fan3d, while
Pollen sub-zone Fan3d: 94–33 cm; ~1670–430 cal. yr BP
Poaceae decreases to 70–90% of the pollen sum, whereas
Pollen sub-zone Fan3e: 33–0 cm; ~430 cal. yr BP to recent
Poaceae percentages increase and
PCA
PCA was used to investigate statistical relationships within the pollen dataset, which could help to identify floral assemblages composed of co-occurring taxa with similar environmental preferences. The first and second PCA axes explain 34% of the total variation. The first PCA axis explains 20% of the variance and clearly separates the dominant taxa

PCA of the Fan Lake pollen data showing the scores for the main pollen types (

Overview figure showing loss-on-ignition (LOI 550°C) (wt% loss), total long-distance taxa:
Confidence intervals of long-distance taxa
The 95% confidence intervals were plotted for the long-distance taxa (Figure 6) to identify significant changes in long-distance pollen transport (a function of SHW strength) and to separate them from random variations inherent in the percentage counts (Maher, 1972). Confidence limits overlap throughout the majority of the Fan Lake record indicating few significant changes in long-distance pollen transport. However, pollen influx of long-distance taxa such as
Discussion
Challenges in the interpretation of sub-Antarctic pollen records
Despite the treeless and phanerogam-poor sub-Antarctic vegetation (Van der Putten et al., 2012b) and the low pollen diversity, the Fan Lake record shows interpretable changes in the major pollen taxa (i.e.
Changes in the relative abundance of pollen from
The Fan Lake record shows a significant change at
Local lake environments
During the mid-Holocene, Fan Lake was surrounded by sub-Antarctic vegetation mainly consisting of small shrubs and herbs, such as
A peak in indeterminable grains, fern and fungal spores at ~3860 cal. yr BP suggests a relatively short disturbance and soil erosion, which coincides with an influx of coarse sand and some of the highest sedimentation rates (0.2–0.4 cm/yr) in the core (Figure 2). The subsequent decrease in the percentage of indeterminable grains at ~3790 cal. yr BP could indicate a slow progression towards a less glaciated mid–upper catchment and may coincide with the onset of a late Holocene ‘climate optimum’. A strong increase in pollen influx and organic carbon suggests higher catchment biomass within lake production between ~3230 and 2710 cal. yr BP. A warmer-than-present environment is inferred from an increase in
From
Two colder/wetter periods can be identified between
Long-distance signal
Long-distance pollen grains from the South American continent are regularly present in the Fan Lake record. Changes in the abundance of the long-distance grains can aid in determining latitudinal shifts or changes in the intensity of the SHW (Bentley et al., 2009; Markgraf et al., 2003; Moreno et al., 2009). The long-distance signal is dominated by pollen from the southern beech
The Fan Lake record shows slight increases in long-distance pollen influx throughout the late Holocene cooling after 2430 cal. yr BP. Small peaks are present between approximately 2210 and 1670 cal. yr BP (Figure 4). The highest abundance of long-distance pollen occurs during the inferred cooler and wetter periods on Annenkov Island.
Climate change
The Fan Lake record provides evidence of warmer conditions in the mid- to late Holocene, which culminates in a period of comparatively higher biomass production between 3790 and 2750 cal. yr BP. With a subsequent gradual change towards slightly cooler and wetter environments after
Previous records from the north coast of South Georgia suggest several cooling and drying events during the mid- to late Holocene, but no strong signal of these changes are present on Annenkov Island. For example, from grey scale density, weight-loss-on-ignition and grain size analyses, Rosqvist and Schuber (2003) reconstructed cooler periods between 7800 and 7400 cal. yr BP, 7200 and 7000 cal. yr BP, 5200 and 4400 cal. yr BP and 2400 and 1600 cal. yr BP. The slight increases in the pollen percentages of the cushion plant
Rosqvist and Schuber (2003) indicate cooling on South Georgia between 2400 and 1600 cal. yr BP based on glacial advances, although the accuracy of these dates have been questioned (Moreton et al., 2004). The interpretation of a late Holocene climate change at Fan Lake is also supported by lake sediment studies on South Georgia (Lake Maiviken), which suggest the onset of wetter conditions at
Various sub- and maritime-Antarctic islands and Antarctic Peninsula records closely resemble Fan Lake in showing a late Holocene ‘climate optimum’, late Holocene cooling and a possible warming overlapping with the ‘Medieval Climate Anomaly’ (‘MCA’) (Table 1). At Fan Lake, the warm and stable lake conditions inferred between 3790 and 2750 cal. yr BP overlap with the late Holocene ‘climate optimum’ from
From 2710 cal. yr BP onwards, the decline in pollen influx in the Fan Lake record indicates a change towards a generally cooler and wetter climate, which persisted until today. With several interbedded warm and cold periods, the climate on Annenkov Island has been very variable throughout the last 2700 years. The majority of research in the sub-Antarctic puts the start of cooling in the late Holocene at
At the end of the late Holocene, increased
SHW and climate changes
The influx of non-native long-distance transported pollen grains in fossil records from the sub-Antarctic islands is primarily controlled by wind strength and the latitudinal position of the SHW and is therefore indicative of changes in atmospheric circulation patterns (e.g. Van der Putten et al., 2012b). Climate models and observational data suggest that during cold periods, the SHW shifts towards the equator, whereas during warmer phases (such as today), the movement is polewards (Bentley et al., 2009; Lamy et al., 2001; Toggweiler et al., 2006; Varma et al., 2012). However, how latitudinal shifts of wind belts affected precipitation and wind strength in the sub-Antarctic region (at
Multi-proxy studies on lake sediment cores in SW Patagonia and Chile show increased wind strength along with increases in precipitation during cooler periods in the late Holocene, such as the ‘Little Ice Age’ (‘LIA’), 400–150 cal. yr BP (Bentley et al., 2009; Moy et al., 2008). Increased wind strength and precipitation associated with the SHW could have caused the expansion of
In contrast, the pollen record from Fan Lake situated at
Conclusion
Most existing palaeoenvironmental records from the Antarctic and sub-Antarctic islands broadly identify a trend from a warm period in the beginning of the late Holocene followed by a prolonged cooling phase, but several uncertainties remain in defining the timing and underlying drivers for these past changes.
The well-dated, 5.8 m-long sediment record from Fan Lake on Annenkov Island, near South Georgia, was used to reconstruct the Holocene vegetation and climate history of the South Atlantic sector of the Southern Ocean at a high temporal resolution. We identified three key phases in the pollen record from Fan Lake: (1) a warm late Holocene ‘climate optimum’ from
The presence of long-distance transport pollen grains in the Fan Lake record was used to reconstruct changes in atmospheric circulation patterns. Although the overall number of long-distance transport pollen grains in the Fan Lake record was comparatively low, the pollen record suggests a link between cold periods and stronger SHW in the late Holocene at this location.
Further studies are needed to better understand the history of the SHW and their impact on Holocene climate change. We suggest that future research should focus on coring sites on the west coasts of sub-Antarctic islands, close enough to the continental landmasses to show a distinct signal of long-distance pollen grain transport.
