Abstract
In the tropical Andes, impacts of both natural and anthropogenic disturbances have been detected over a period exceeding 4000 years. However, the history of the environment remains unknown in most Andean regions. To infer possible interactions between climate and humans, we analysed the pollen content of an 8.5 m deep peat core extracted from a peat bog located near the Nevado Coropuna volcano on the slope of the Western Cordillera in southern Peru (15°30S, 72°40W, 6380 m). Results showed that taxa of the upper Puna expanded when cooler and moister climatic conditions prevailed during a time period that includes the early and the mid Holocene (9700–5200 cal. yr BP). An increase in shrub pollen frequencies and a decrease in the Poaceae/Asteraceae ratio are attributed to a drier climate during the late Holocene (5200–3000 cal. yr BP). After 3000 cal. yr BP, the vegetation cover resembled that of today. Both archaeological and pollen data attest to the beginning of agriculture from 2200 cal. yr BP. Around 900 cal. yr BP, the vegetation cover suddenly changed, probably because of a colder and drier climate. Past societies continued their agricultural activities despite this abrupt change. Our results emphasize the specific geographical location of Nevado Coropuna astride the Western Cordillera and the western edge of the Altiplano, which is consequently subject to both Pacific and Atlantic influences.
Introduction
Long-term climate records are necessary to reveal the influence of climate changes on human societies and of man’s impact on the environment (Brenner et al., 2001; Hodell et al., 1995; Weiss and Bradley, 2001), to increase our knowledge of global climate changes and to calibrate numerical simulations of global circulation (Markgraf et al., 2002; Messerli et al., 1993; Smol and Cumming, 2000; Sterken et al., 2006; Thompson et al., 1998). In the tropical Andes, both climate and humans have impacted the landscape. Several human communities, including the Inca Empire, occupied the tropical Andes for thousands of years while changes in climate and vegetation occurred under the influence of both Pacific and Atlantic circulations. In recent decades, palaeoenvironmental studies have been carried out from southern Ecuador to northern Chile in the Central Andes revealing contrasted situations depending on the latitude (e.g. Abbott et al., 2003). For instance, during the mid Holocene, between 8000 and 5000 cal. yr BP, the region of Lake Titicaca was characterized by a dry episode with the lowest water level recorded in 30 000 years (Abbott et al., 1997a; Baker et al., 2001; Cross et al., 2001; Hanselman et al., 2005; Mourguiart, 1990; Paduano et al., 2003; Seltzer et al., 1998; Ybert, 1988). New radiocarbon dates obtained from recently exposed plant remnants at high elevations (Nevado Quelccaya, eastern Cordillera, southern Peru) also attest to a minimal extension of the glacier at 5200 cal. yr BP (Buffen et al., 2009). In contrast, in northern Chile, packrat middens and geomorphological analyses attest to a more humid episode during the mid Holocene in what is today the Atacama Desert (Betancourt et al., 2000; Quade et al., 2001). Strong latitudinal contrasts may explain these regional differences based on modern settings and observations of the different atmospheric circulation patterns between the Altiplano and the Atacama region characterising a transition area with either an Atlantic or a Pacific influence on the yearly distribution of precipitation (Vuille and Keimig, 2004). Another abrupt and drastic dry episode was recorded at 850 cal. yr BP at different locations in the Central Andes, e.g. Lake Titicaca (Abbott et al., 1997b; Binford et al., 1997) and Nevado Quelccaya (Ortloff and Kolata, 1993; Thompson et al., 1985). This event has been linked with the disappearance of the Tiwanaku civilization (Binford et al., 1997).
However, since their original expansion, economic activities and urban centres have mainly been located on the Pacific coast and in the western Cordillera where the climate is arid, with less than 250 mm precipitation per year. The development of human communities and civilizations that lasted for centuries in these dry areas was possible because man adapted to his environment and water was available from melting glaciers (Chepstow-Lusty et al., 2009). Detailed knowledge on related changes in climate and societies at high elevations is still needed to evaluate the future of some urban centres in a scenario without glaciers and given the predicted changes in the distribution of precipitation (Bradley et al., 2006; Urrutia and Vuille, 2009). To improve the current knowledge on climate and environmental changes in the western Cordillera, we present a new pollen record located in an inhabited highland near the Nevado Coropuna in southwestern Peru. Our interpretation of fossil pollen data is supported by published modern pollen rain results based on 24 samples collected from different types of vegetation (Kuentz et al., 2007). We combined archaeological data with pollen analysis to evaluate the past impact and the adaptation of human activities to arid landscapes.
Modern setting: Climate and vegetation in southern Peru
The Nevado Coropuna (15°30S, 72°40W) is one of the Quaternary volcanoes that form the Peruvian volcanic arc in the Central Andes. It straddles the western Cordillera and the western edge of the Altiplano at an altitude of 6380 m a.s.l. The western Cordillera is characterized by steep temperature and precipitation gradients over a distance of only 120 km between the arid Pacific coastline and the snowline (now c. 5400 m a.s.l.) at latitude 15°S.
Today, variations in insolation between austral winter and summer are high and are responsible for shifts in the Intertropical Convergence Zone (ITCZ). The southern position of the ITCZ allows penetration of humid air masses from the Atlantic favored by the easterly wind circulation termed South American Summer Monsoon SASM (Vuille and Keimig, 2004; Zhou and Lau, 1998) when summer precipitation occurs on the Altiplano and around Nevado Coropuna. About 70–80% of total precipitation falls in summer. On the other hand, a dominance of the westerly wind circulation can favor the installation of dry conditions in our study area. Seasonal and interannual variability is considerable because of an increase/decrease in the surface temperature of the Pacific Ocean during El Niño/La Niña phases of the Southern Oscillation (ENSO). Strong El Niño events such as the one in 1998 can cause dramatic flooding along the northern Peruvian coast (Fontugne et al., 1999), and retreat of mountain glaciers in northern Bolivia (Francou et al., 2003) and in southern Peru (Forget et al., 2008; Thouret et al., 2002). Prolonged La Niña-like conditions are induced by cool ocean surface temperature anomalies in the central and eastern tropical Pacific (Trouet et al., 2009), and can lead to glacier expansion (Arnaud et al., 2001; Francou et al., 2004).
The main vegetation type in the tropical Andes is high elevation grassland also called Puna, which covers an area of c. 85 000 km2. Today, Puna grows between 3500 and 5200 m a.s.l., between the steppe and the snowline in the western Cordillera of Peru, and between the Andean rain forest and the snowline in the eastern Cordillera of Peru (Kuentz et al., 2007) (Figure 1A). Five altitudinal vegetation types were identified around Nevado Coropuna and are described in Kuentz et al. (2007) and Kuentz (2009) (Table 1 and Figure 1B). These vegetation types include, from the lowest to the highest in elevation: (1) The Ambrosia steppe (800–2500 m a.s.l.) is mostly composed of Ambrosia shrubs. (2) In the pre-Puna (2500–3500 m a.s.l.), the vegetation cover is denser and includes different shrubs of the Asteraceae family (Ambrosia, Diplostephium, Senecio). (3) The lower Puna (3500–4000 m a.s.l.) is composed of Asteraceae shrubs and herbaceous species of the Fabaceae and Solanaceae families. (4) In the Puna sensu stricto (4000–4800 m a.s.l.) the grassland is mainly composed of grasses (e.g. Calamagrostis, Festuca, Stipa) and shrubs of the Asteraceae family (Baccharis, Chuquiraga, Parastrephia, Senecio) (UNESCO, 1981). Patches of Polylepis forest grow around 4400 m a.s.l. The flat bottoms of glacially shaped valleys are often occupied by peat bogs (Cyperaceae, Juncaceae). (5) In the upper Puna (4800–5200 m a.s.l.), the sparse vegetation is composed of Poaceae and Asteraceae shrubs (Baccharis, Senecio, Wermeria). Cushions of Apiaceae and Caryophyllaceae families are abundant around 5000 m a.s.l.
Vegetation types around Nevado Coropuna with their corresponding pollen indicators and climatic parameters (from Kuentz et al., 2007)

The biogeographical and palaeoenvironmental setting of the Nevado Coropuna volcano. (A) Location of Nevado Coropuna in the study area, main vegetation belts in Peru, and palaeoenvironmental record sites quoted in this study. (B) Map of the vegetation types around Nevado Coropuna with the location of the archaeological sites. Distinct symbols are keyed to the archaeological chronological frame. Locations of sampled cores are also shown
Materials and methods
Vegetation types were mapped by remote sensing using a 2002 Landsat ETM image (Figure 1B) (Kuentz et al., 2007). A total of 24 soil-surface samples of all vegetation types were collected. In 1998, an 8.5 m deep sedimentary core (COR300) was drilled with a Russian corer in a peat bog located at 4400 m a.s.l. in a large glacially shaped valley that drains the SSW of Nevado Coropuna (Figure 1B). This core was extruded in a vertical position to maintain stratigraphy, sediment samples were wrapped in the field and shipped unopened in their polycarbonate coring tubes to the University of Montpellier, France. The core (COR300) was analysed with an average sampling resolution of 6 cm. The first 150 cm cover the last millennium and analyses are still in progress. Pollen grains were extracted from the sediments following the standard protocol developed by Fægri and Iversen (1989). The peat is preserved because of slow decomposition under saturated conditions; therefore, significant bioturbation is unlikely. Pollen concentrations were calculated according to the method of Cour (1974). Pollen counts were performed under 600× magnification and morphology was identified by comparison with our reference pollen collection obtained from floristic materials collected in the field and from herbariums (Universidad Nacional San Augustin de Arequipa, Museo de Historia Natural de Lima and Herbario Nacional de Bolivia), as well as by comparison with pollen atlases (Heusser, 1971; Hooghiemstra, 1984). More than 300 pollen grains were counted per sample. The total pollen sum included arboreal and non-arboreal pollen, but excluding Poaceae, Cyperaceae and Juncaceae, which are over-represented with respect to the frequencies of other taxa. Influx of charcoal particles was estimated according to Clark (1982). This is a point counting method to estimate the projected area of charcoal. The pollen diagram was arranged according to results of modern pollen rain analyses. The taxa were organized in accordance with the vegetation strata (trees/shrubs/herbs) except for the peat-bog taxa (local production). These results of the modern pollen rain revealed the presence of pollen taxa at different altitudinal ranges and an altitudinal pollen gradient was deduced: Asteraceae type Ambrosia (1800–2200 m a.s.l.), Malvaceae (2700–3300 m a.s.l.), Asteraceae-type Senecio (3500–4100 m a.s.l.) and Apiaceae (above 4600 m a.s.l.) (Kuentz et al., 2007). Concerning fossil data, an ecological diagram was built to reflect this altitudinal pollen gradient. We extrapolated the precipitation range around Nevado Coropuna based on meteorological data from SENAMHI (Servicio Nacional de Meteorologia e Hidrologia). The Poaceae/Asteraceae (P/A) ratio calculated for our pollen surface samples was plotted and arranged according to a precipitation gradient (Figure 2). Except for one sample (25), the P/A ratio is well correlated with changes in precipitation, in agreement with the humidity index defined by Reese and Liu (2005), and can thus be used to interpret the fossil data. Constrained clustering was performed using Zone software (Juggins, 2005). AMS 14C datings of seven peat samples was carried out at the Laboratoire de Mesure du Carbone (Gif sur Yvette, France) in 2007. Calibrated ages were obtained with Calib501 software (Stuiver et al., 2005) (Table 2). We applied a Southern Hemisphere correction to all samples (McCormac et al., 2004). Data on human occupation during the Holocene was acquired from previous archaeological and geographical studies (Forget et al., 2008; Ziolkowski, 2005; Ziolkowski and Belán Franco, 2001) and georeferenced in a Geographic Information System.

(A) Map showing the extrapolation of the precipitation range around Nevado Coropuna based on data from SENAMHI (Servicio Nacional de Meteorologia e Hidrologia) with the location of the pollen surface samples (from Kuentz et al., 2007). (B) Poaceae/Asteraceae ratio of the soil-surface samples
Radiocarbon dates for the COR 300 peat core with calibrated ages using a Southern Hemispheric correction according to CALIB 5.0.1 (Stuiver et al., 2005)
Present human and archaeological data
Present human activities are described as a function of the altitudinal vegetation types found from low to high elevation. The Ambrosia steppe is irrigated in valley floors and used for growing onions, sugar cane, potatoes, and rice. In the pre-Puna, vegetables, maize and fodder crops are the main crops. Pasture is also extensive in this vegetation type. Fuel wood is provided by planting eucalyptus and pines. In the lower Puna, dry agriculture with tubercles (Oxalis, Solanum), Quinoa and Kiwicha predominate. In the Puna sensu stricto, when the ground used for livestock breeding becomes dry and compacted by animals such as lamas and alpacas, the Poaceae Stippa ichu predominates. When livestock graze humid soils, Pycnophyllum molle prevails. In this vegetation type, villages generally depend on glacier meltwater. Polylepis forests are frequently harvested for fuel as they are the only source of wood at this elevation. Medicinal plants have also been identified in this vegetation type (Brack Egg, 1999). In the upper Puna, the impact of human activity is limited to temporary camps and religious monuments.
Numerous archaeological sites located in southern coastal Peru provided evidence for a first occupation by hunter-gatherers between 13 000 and 11 000 cal. yr BP (Sandweiss, 2003; Sandweiss et al., 1998). High elevation areas are thought to have been colonized after c. 11 000 yr BP as evidenced by ancient paths (Aldenderfer, 1999; Baied and Wheeler, 1993; Marchant et al., 2004). The domestication of the camel family in the Andes dates back to 6000 yr BP (Guffroy, 1999; Jakubicka and Woloszyn, 2005). At Nevado Coropuna, two caves occupied by hunter-gatherers have been identified and dated before 5000 cal. yr BP (Figure 1B). No archaeological artefacts have been found for the period between 3000 and 2300 cal. yr BP around Coropuna. Between 2300 and 1000 cal. yr BP, four archaeological sites have been identified in the region of Coropuna (Figure 1B), all located in the pre-Puna vegetation type at 3000 m a.s.l. As many as 30 archaeological sites dated between 1000 and 700 cal. yr BP are located in the pre-Puna, lower Puna and Puna sensu stricto. They attest to the spread and diversification of pastoral and agricultural activities between 2500 and 4800 m a.s.l. at that time. During the Inca Empire (
Results
Chronology and stratigraphy
The core was composed of brown peat with four thin layers of non-organic material (silt and/or clay). The chronology of COR300 core is based on seven AMS radiocarbon dates from bulk. As no age reversal was observed, these dates were all used to reconstruct a timescale. The age model was drawn with Kaleidagraph software from a smooth curve (Figure 3). The oldest age, at the base of the core, was estimated to be 9750 cal. yr BP. Sedimentation rates varied over time. From 9750 to 6600 cal. yr BP, the average rate of sedimentation was 0.5 mm/yr, increased between 6600 and 4700 cal. yr BP (1.7 mm/yr on average), but decreased between 4700 and 1500 cal. yr BP, it decreased (0.7 mm/yr on average). Between 1500 and 800 cal. yr BP, the sedimentation rate was very high (3.7 mm/yr on average).

Radiocarbon dates used to build the age model and changes in sedimentation rates along the core COR 300
Pollen zones
Cluster analysis was performed on the pollen counts to define pollen zones (Figure 4). However, our results did not enable us to draw any subdivisions in the diagram probably because all taxa have the same statistical weight in the cluster. Consequently, we based our pollen zonation on a comparison with modern pollen rain results and used visible changes in the indicator taxa (mainly Ambrosia, Apiaceae, Poaceae, the Poaceae/Asteraceae ratio and pollen concentration). We were able to characterize four pollen zones (Figure 4). Using light microscopy, pollen analysts are unable to distinguish Acaena from Polylepis. Acaena is currently present in warm humid valleys and can be confused with Polylepis in the eastern Cordillera (Chepstow-Lusty et al., 2005). Because Acaena was not observed during our vegetation surveys and has never been recorded by botanists at the Universidad Nacional San Augustin de Arequipa in the study area, we refer only to Polylepis pollen grains.

Palynological diagram of core COR 300, taken 8 km SSW of the summit of the Nevado Coropuna volcano (Southern Peru). Pollen frequencies, concentrations and influx, charcoal influx, and a summary ecological diagram are presented along a depth scale. The dotted line indicates the beginning of agriculture around Nevado Coropuna
Zone 1. Early and mid Holocene (996–691 cm, 9700–5200 cal. yr BP, 51 samples)
Zone 1 was characterized by high frequencies of Apiaceae (9.5%), Brassicaceae (3.7%), Poaceae (85%) and Polylepis (5.2%) and relatively low frequencies of Ambrosia (16.6%). Charcoal influx was relatively high (1.2 cm2/cm3 per yr). The rate of sediment accumulation was very low (0.3 mm/yr) in the lower part of the zone, but increased to 1.4 mm/yr at a depth of 840 cm (c. 6300 cal. yr BP). Concentrations were relatively high in the lowest part of this zone and reached maximum at a depth of 930 cm, i.e. c. 7500 cal. yr BP with 12 600 grains/g.
Zone 2. Late Holocene 1 (691–481 cm, 5200–3000 cal. yr BP, 36 samples)
This zone was characterized by very low pollen concentrations (363 grains/g) and a decrease in Apiaceae (6%) and Poaceae (72%) frequencies. Ambrosia (17.3%), Ophryosporus (6%) and Chenopodiaceae/Amaranthaceae (12%) frequencies slightly increased. Polylepis frequencies were low (4.1%) and decreased drastically (2%) after c. 4600 cal. yr BP (at 571 cm in depth). At the same time, Apiaceae and Brassicaceae frequencies also decreased (3% and 2%, respectively). Charcoal particles were rare in this zone (influx of 0.5 cm2/cm3 per yr). Sediment accumulation accelerated from 1.4 mm/yr to 2.2 mm/yr (at 613 depth, i.e. c. 4900 cal. yr BP), before dropping to 0.5 mm/yr at c. 4000 cal. yr BP.
Zone 3. Late Holocene 2 (481–199 cm, 3000–900 cal. yr BP, 43 samples)
This zone contained very high frequencies of Ambrosia (26%). Frequencies of Poaceae (82%) and concentrations values (1030 grains/g) increased in comparison with the previous zone. Ophryosporus (10%) and Chenopodiaceae/Amaranthaceae (15%) frequencies increased slightly whereas frequencies of Apiaceae (5%) and Polylepis (2.2%) remained low. Few charcoal particles were counted in this zone (influx of 0.4 cm2/cm3 per yr). The rate of sediment accumulation was high (2.3 mm/yr on average). A break in pollen spectra was identified at c. 2200 cal. yr BP (at a depth of 433 cm). After this date, frequencies of Ambrosia increased (from 22 to 26%) while frequencies of Apiaceae decreased (from 6 to 4%). Solanaceae were occasionally identified from time to time (0.3%) whereas they were uncommon before c. 2200 cal. yr BP (Figure 5).

Synthetic palynological diagram from core COR 300 (only the taxa cited in the discussion are shown). Pollen frequencies, concentrations, and charcoal influx are presented along a time scale. P1, P2, P3 and P4 are the pollen zones. The dot indicates the domestication of the camelideae in the Andes. The star indicates the beginning of agriculture around Nevado Coropuna
Zone 4. Recent period (199–157 cm, about 900–800 cal. yr BP, 8 samples)
This zone was characterized by a marked decrease in the frequencies of Poaceae (50%), Ambrosia (18%), Ophryosporus (5%), Chenopodiaceae/Amaranthaceae (4%) and an increase in Apiaceae (10%), Caryophyllaceae (17%) and Brassicaceae (4%) frequencies. Pollen concentrations were very low (500 grains/g). Arboreal pollen taxa Polylepis almost disappeared. Pollen grains of Rosaceae (0.4%) were identified for the first time, and Solanaceae (0.3%) were still present. Malvaceae (1%), Fabaceae (1.9%) and Alnus reached their highest frequencies. The influx of charcoal particles remained low (0.3 cm2/cm3 per yr). This zone showed the highest rate of sedimentation (4.1 mm/yr on average).
Discussion
The early–mid Holocene (9700–5200 cal. yr BP): A moist and cold climate
Between 9700 and 5200 cal. yr BP, Apiaceae cushions and Polylepis forest expanded in Nevado Coropuna, whereas Ambrosia shrubs are rarely recorded (Figure 5). According to pollen rain analysis, this assemblage matches a well-developed upper Puna belt and suggests moister and colder conditions than today (Table 1) (Kuentz et al., 2007). However, in the other Andean palaeoenvironmental records, this time interval is generally divided into two distinct climatic periods. The first period of time between c. 10 000 and 7500 cal. yr BP, represents the early Holocene and the second, between 7500 and 5000 cal. yr BP, characterizes the mid Holocene. During the early Holocene, 120 km northwest of Nevado Coropuna, data based on snail shells and loess attest to moister conditions at c. 9000 cal. yr BP (Unkel et al., 2007). Palynological and sedimentological analyses show that 250 km to the east, the level of Lake Titicaca was relatively high between 10 000 and 8500 cal. yr BP (Baker et al., 2001; Paduano et al., 2003). Geomorphological studies on moraines formed during glacial stages attest to the fact that one glacier in northern Bolivia reached low elevations between 10 000 and 7000 cal. yr BP (Smith et al., 2011). Further south, archeology and lake level analyses show similar moist conditions between 12 900 and 8400 cal. yr BP in the Atacama region (22–24°S) (Grosjean et al., 2001; Messerli et al., 1993; Wolfe et al., 2001). The slightly moister and colder conditions deduced in the Nevado Coropuna record could be linked with prevailing La Niña-like conditions resulting in an increase in precipitation on the Altiplano (Arnaud et al., 2001; Francou et al., 2004). La Niña-like conditions are generally linked with colder/warmer tropical Pacific Ocean surface temperatures in the eastern/western parts of the Ocean than today, associated with weaker and less frequent El Niño events (Shin et al., 2006). Various proxy reconstructions inferred such climatic conditions during the early Holocene (Overpeck and Webb, 2000; Sandweiss et al., 2001; Tudhope et al., 2001).
From c. 8000 cal. yr BP, the climate became progressively drier in the southern Tropics and moister in the northern Tropics (Haug et al., 2001). However, close to Nevado Coropuna and at lower elevations, the level of Lake Pacucha (c. 300 km to the northeast) was lower during this interval, but the vegetation remained abundant, attesting to high seasonal moisture rates (Hillyer et al., 2009; Valencia et al., 2010). Conversely palaeo-proxies from Lake Titicaca records attest to extreme aridity characterized by a sharp decrease in the lake level to as much as 100 m below its present level (e.g. Argollo and Mourguiart, 2000; Baker et al., 2001; Hanselman et al., 2005). Geochemical analyses also revealed an arid period between 8600 and 6400 cal. yr BP at Lake Chungará located at the border between Chile and Bolivia (Moreno et al., 2007). Further south (23°S), the record of Laguna Miscanti also reflects an arid climate during the mid Holocene although frequent changes in moisture rates are attested (Grosjean et al., 2001). A northernmost position of the Intertropical Convergence Zone at millennial-scale is observed either by proxy data or by simulations during the mid Holocene (Braconnot et al., 2007). This situation prevented the summer rainy season in the southern Tropics and produced a dry climate similar to that recorded in the Altiplano. Further south, in the Atacama region, the situation was different. Analyses performed on lake terraces around Lake Aricota (17°S), U/Th analysis of the sediment from the Salar of Atacama (23°S) and fossil rodent middens recorded in the central Atacama (22–24°S) revealed a moister environment during the mid Holocene than at present (Betancourt et al., 2000; Bobst et al., 2001; Latorre et al., 2005; Placzek and Quade, 2001; Placzek et al., 2006). The interpretation of drier mid-Holocene conditions around the Titicaca basin and conversely moister conditions in the Atacama region has been the subject of debate in the scientific community (Betancourt et al., 2000; Grosjean, 2001; Quade et al., 2001). In reality, the different proxies used by these authors (packrat-middens versus lake levels) recorded climate variability at various timescales: consequently the mid-Holocene period was generally dry but included humid intervals at annual and decadal scales (Grosjean et al., 2003). The specific location of Nevado Coropuna in the transition area between the Altiplano and the Pacific slope of the Central Andes could explain why no break in the vegetation cover was observed between the early and the mid Holocene, in contrast to other records in the Central Andes. During the early Holocene, the Titicaca basin, the Atacama region and Nevado Coropuna were correlated, showing similar moister conditions. In contrast, during the mid Holocene, climatic dynamics may have differed between the Altiplano and the western slopes of the Andes. During this period, the Nevado Coropuna record was similar to records of the Atacama region but differed from records of the Altiplano. These conclusions support the hypothesis of a highly variable precipitation pattern over the Central Andes (Vuille and Keimig, 2004; Vuille et al., 2000).
Mid to late Holocene: Installation of drier climatic conditions
Between 5200 and 3000 cal. yr BP, the vegetation around Nevado Coropuna was made up of a majority of shrubs of the Puna sensu stricto. After the early Holocene, the herbaceous vegetation (majority of Poaceae) decreased in favour of the shrub vegetation (Asteraceae mainly). The Poaceae/Asteraceae (P/A) ratio and the pollen concentrations were low between 5200 and 3000 cal. yr BP, attesting to dry climatic conditions (Figures 2 and 6). During this dry interval, the size of the Polylepis forests decreased considerably. For this period, only two caves probably occupied by hunter-gatherers have been identified in the region. Although agriculture started at least as early as 4000 cal yr BP in southern Peru as attested to for instance at Waynuna (Perry et al., 2006), human presence around Coropuna was rare at this time and we consequently excluded any major human impact such as deforestation or terraces constructed for cultivation purposes. At the same latitude as Coropuna but 150 km further east, the Titicaca basin showed arid conditions followed by a slow rise in the lake level after 5000 cal yr BP (Abbott et al., 2003) (Figure 6). On the Nevado Sajama (400 km southeast of the Nevado Coropuna), between 5000 and 3400 cal. yr BP, high dust concentrations in the ice core and low snow accumulation suggest drier conditions (Thompson et al., 1998) (Figure 6). At the end of this interval, around 3000 cal. yr BP, there was an increase in the P/A ratio and in pollen concentrations in the Coropuna pollen record: this suggests the return of moister climatic conditions in the central Andes, in agreement with the ice core record of Nevado Sajama. We interpret these climate changes as being linked with precession cycles and with the southward displacement of ITCZ, which influenced precipitation on the Altiplano (Placzek et al., 2006). Between c. 5000 and 3000 cal. yr BP, the increase of the summer insolation allowed a broader-scale effect of the upper-air easterlies that brought summer precipitation to the Titicaca basin. However we infer that it was only from 3000 cal. yr BP that the intensity of the upper-air easterlies was high enough to allow a weakening of the westerlies that led to a summer precipitation climate regime on Nevados Sajama and Coropuna, as described by Vuille and Keimig (2004) (Figure 6). In addition, the increase in El Niño frequencies observed during this period could also have led to the dry conditions observed at Sajama and Coropuna (Maldonado and Villagrán, 2006; Moy et al., 2002; Riedinger et al., 2002; Rodbell et al., 1999). In conclusion, between 5200 and 3000 cal. yr BP, the human impact alone cannot explain the observed changes in vegetation cover and, here again, the influence of climate prevails.

Comparison between proxy data for palaeo-precipitations in the Central Andes and results of palynological analysis of core COR 300. (A) Fluctuations in the level of Lake Titicaca, as inferred from δ13C measurements (Abbott et al., 2003). (B) Number of particles >0.63 µm/ml and snow accumulation in the Nevado Sajama ice core (Thompson et al., 1998). (C) Poaceae/Asteraceae (P/A) ratio and pollen concentration (Cn) in COR 300 core. (D) January insolation at 15°S (Berger, 1992)
Human impact on vegetation
Between 2200 and 900 cal. yr BP, the taxa indicator of the upper Puna (mainly Apiaceae) decreased while the herbaceous and shrub vegetation (mainly Ambrosia) indicator of the lower Puna increased (Figure 5). This pollen assemblage characterizes similar climatic conditions as today. The expansion of Ambrosia and the regression of Apiaceae could also reflect an increase in temperature and/or drier climatic conditions (Table 1) (Kuentz et al., 2007). One possibility is that these changes were linked with the increase in El Niño frequencies that characterized this time period (Maldonado and Villagrán, 2006; Moy et al., 2002; Riedinger et al., 2002). However, we know that an increase in ENSO frequencies could not have impacted the climate of the Central Andes because neither Lake Titicaca nor the Sajama ice core recorded drier conditions during this period (Figure 6). Around the Nevado Coropuna, climatic conditions were relatively humid with summer precipitation as revealed by the increase in the P/A ratio (Figure 6). A more probable hypothesis is that changes in the vegetation cover were caused by human activities. Indeed, around the Nevado Coropuna, four archaeological sites date back to the Intermediate Period (2300–1000 cal. yr BP) at 3000 m a.s.l., and are linked with the first agricultural activities in the region (Figure 1B) (Kuentz, 2009). The increase in Ambrosia could also reflect the plantation of these shrubs to stabilize agricultural terraces (Chepstow-Lusty et al., 1998). The increase in Solanaceae and Chenopodiaceae/Amaranthaceae, which are native cultivated taxa, also supports this hypothesis (Figure 5). In addition, the fact that Azorella compacta was one of the main sources of fuel for human populations may explain the regression of Apiaceae cushions in the area. Both archaeological and pollen data attest to high human presence and to the beginning of agriculture from 2200 cal. yr BP in the region of Coropuna. Therefore changes in cover vegetation during this period should be attributed to past societies and less to climatic changes.
An abrupt and short environmental change (ad 970–1130)
Between 980 and 820 cal. yr BP (about
Charcoal record of the core COR300
Because fires were rare in the Amazon basin between 11 000 and 7500 cal. yr BP (Power et al., 2008), the source of the charcoal particles observed in core COR 300 near Nevado Coropuna during this period could be linked with local storms and/or volcanism. On the other hand, between 8000 and 4000 cal. yr BP, fires increased in the Amazon basin and on the eastern slopes of the Andes (Mayle and Power, 2007). Therefore, after 8000 cal. yr BP, the charcoal particles observed in the sediment around Nevado Coropuna may have been carried aloft from their area of origin in the south and southeast.
Despite increasing fires in tropical South America around 3000 cal. yr BP (Power et al., 2008), the frequency of charcoal particles remained low at Nevado Coropuna after the mid Holocene. Our results show that the amount of biomass that could be used as fuel in these arid landscapes was reduced, with rare Polylepis trees and Azorella compacta cushions, thus preventing the occurrence of big fires. However, the presence of small local fires cannot be excluded, either man-induced or natural (e.g. volcanic eruptions in the Andahua field of monogenetic volcanoes 25–30 km east of Nevado Coropuna or thunderstorms).
Conclusion
The ‘Nevado Coropuna’ record is the first pollen record located in the transition area between the western Cordillera and the Altiplano in southern Peru, and which covers the whole Holocene. Our research shows that major environmental changes occurred throughout the Holocene even in the dry vegetation cover at high elevations in the Central Andes. These changes were linked with climatic variability of the Central Andes distinguishing and alternating the influence of Pacific or Atlantic circulations, which is in agreement with modern observations and model predictions for the twenty-first century in the Central Andes (Urrutia and Vuille, 2009; Vuille et al., 2000). The moist conditions that prevailed during the early and mid Holocene (9700–5200 cal. yr BP) were correlated with prolonged La Niña-like conditions and cooler surface temperatures of the Pacific Ocean (Carré et al., 2011), whereas the drier late-Holocene (5200–3000 cal. yr BP) climate was attributed to a weaker influence of the westerlies. Agriculture appears to have begun later in this region than in other parts of the Central Andes but the succession of late-Holocene and historical civilizations strongly impacted the vegetation cover from 2200 cal. yr BP onward. Between 980 and 820 cal. yr BP an abrupt environmental change was probably due to colder and drier conditions although it did not affect agricultural activities. In the twenty-first century, climate simulations (e.g. Intergovernmental Panel on Climate Change (IPCC), 2007) predict an increase in aridity in the southern tropical Andes and/or an increase in rainfall in the northern tropical Andes (Solomon et al., 2009; Urrutia and Vuille, 2009). Many tropical glaciers will almost completely melt in the next 50 years and the continuous increase in the population will lead to more livestock grazing in the highlands (Bradley et al., 2006).The contrasted patterns observed at Coropuna during the Holocene confirm the fragility of these high-elevation environments in the face of a changing climate and reveal the importance of geographical location in evaluating the consequences of probable changes in the lower tropical belt (Seidel et al., 2008). These observations underline the need for more environmental studies in the tropical Andean highlands if appropriate strategies are to be found to enable the populations to survive in a drying landscape, because water resources usually used for irrigation will become scarce in the near future.
Footnotes
Acknowledgements
Radiocarbon dating was performed at the Laboratoire de Mesure du Carbone 14 (LMC14) – UMS 2572 (CEA/DSM – CNRS – IRD – IRSN – Ministère de la culture et de la communication) at Gif sur Yvette, France. This research is part of the IRD (Institut de Recherche pour le Développement) ‘GREAT ICE’ programme and was funded by IFEA (Institut Français des Etudes Andines), IRD and the CERAMAC laboratory (University Clermont-Ferrand 2). AK thanks Denis Didier Rousseau, Rachid Cheddadi and the ‘Paléoenvironnements et Paléoclimats’ team of ISEM, Stephan Beck and the National Herbarium at La Paz (Bolivia) for welcoming her during her PhD. Thanks to an anonymous reviewer who has significantly improved the manuscript.
