Earth-Science Reviews 97 (2009) 242–256 Contents lists available at ScienceDirect Earth-Science Reviews j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / e a r s c i r ev Vegetation response to Holocene climate change in monsoon-influenced region of China Yan Zhao a,⁎, Zicheng Yu b, Fahu Chen a,⁎, Jiawu Zhang a, Bao Yang c a b c MOE Key Laboratory of Western China's Environmental System, College of Earth and Environmental Sciences, Lanzhou University, Lanzhou 730000, China Department of Earth and Environmental Sciences, Lehigh University, Bethlehem, PA 18015, USA Cold and Arid Regions Environment and Engineering Research Institute, Chinese Academy of Sciences, Lanzhou 730000, China a r t i c l e i n f o Article history: Received 26 April 2009 Accepted 16 October 2009 Available online 26 October 2009 Keywords: Holocene vegetation change climate change monsoonal China fossil pollen human activity a b s t r a c t Fossil pollen records from 31 sites with reliable chronologies and high-resolution data in the monsoonal region of China were synthesized to document Holocene vegetation and climate change and to understand the large-scale controls on these changes. The reconstruction of moisture histories was based on a four-class ordinal wetness index at 200-year time slices at individual sites. The vegetation experienced diverse changes over the Holocene in different regions: (1) between tropical seasonal rain forest and more open forest in tropical seasonal rain forest region; (2) from mixed evergreen and deciduous broadleaved forest to more deciduous or Pinus-dominated forest in subtropical region; (3) from mixed evergreen and deciduous broadleaved forest to deciduous forest in temperate deciduous forest region; (4) from deciduous broadleaved forest to conifer–deciduous forest in conifer–deciduous mixed forest region; (5) from steppe forest to steppe in temperate steppe region; and (6) from steppe forest/meadow to meadow/steppe in highland meadow/steppe region. Despite various vegetation sequences in different regions, our synthesis results show that a humid climate generally characterized the early and middle Holocene, and a drier climate prevailed during the late Holocene, with an abrupt shift at ca. 4.5 ka (1 ka = 1000 cal yr BP). Abrupt palynological changes based on a squared-chord distance of pollen assemblages occurred at 11–10, 6–5 and 2–1 ka from most sites. The synthesized pattern of moisture change is similar to the ones inferred from other independent climate proxies; however, gradual vegetation changes in the early Holocene lagged about 1000 yr behind the summer monsoon maximum as indicated by speleothem isotope records from Dongge and Sanbao caves. Human activities likely affected vegetation change greatly during the late Holocene, but the magnitude and precise timing are less clear and require further investigation. © 2009 Elsevier B.V. All rights reserved. Contents 1. 2. 3. 4. 5. Introduction . . . . . . . . . . . . . . . . . . . . . . . Data sources and methods . . . . . . . . . . . . . . . . 2.1. Study region and site selection . . . . . . . . . . . 2.2. Moisture index and statistical analysis . . . . . . . Pollen records of Holocene vegetation and climate changes . 3.1. Tropical monsoonal rain forest . . . . . . . . . . . 3.2. Subtropical evergreen and deciduous forest . . . . . 3.3. Temperate deciduous forest . . . . . . . . . . . . 3.4. Temperate mixed conifer–hardwood forest . . . . . 3.5. Temperate steppe . . . . . . . . . . . . . . . . . 3.6. Highland meadow and steppe . . . . . . . . . . . Discussion . . . . . . . . . . . . . . . . . . . . . . . . 4.1. Temporal and spatial patterns of Holocene vegetation 4.2. Vegetation responses to climate oscillations . . . . . 4.3. Human disturbance during the late Holocene . . . . Concluding remarks. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . and . . . . . . ⁎ Corresponding author. Tel.: +86 931 891 2337; fax: +86 931 891 2330. E-mail addresses: [email protected] (Y. Zhao), [email protected] (F. Chen). 0012-8252/$ – see front matter © 2009 Elsevier B.V. All rights reserved. doi:10.1016/j.earscirev.2009.10.007 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . climate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . change. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 243 243 243 244 246 246 246 248 248 248 249 249 249 252 254 255 Y. Zhao et al. / Earth-Science Reviews 97 (2009) 242–256 Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. Introduction The East Asian monsoon system is among the most dynamic continent–ocean–atmosphere systems in the world. Emerging evidence suggest that the summer monsoon intensity followed the summer insolation trend, with the maximum monsoon occurring in the early Holocene and decreasing afterwards (Kutzbach, 1981; COHMAP, 1988; Morrill et al., 2003; Ruddiman, 2008). This trend has been documented in various proxy records in monsoonal China, including speleothem isotopic records from Dongge Cave (Yuan et al., 2004; Dykoski et al., 2005; Wang et al., 2005), Sanbao Cave (Shao et al., 2006; Wang et al., 2008) and Heshang Cave (Hu et al., 2008). Other attempts were also made to study the Holocene climate pattern in monsoon-influenced region of China. Shi et al. (1992) proposed that Holocene Megathermal period happened at 8500–3000 14C yr BP (ca. 9.4–3.1 ka; 1 ka = 1000 cal yr BP) on the basis of pollen data, fossil fauna, paleosol, lake level, glacial remains, and archaeological data. An et al. (2000) proposed that the Holocene East Asian summer monsoon precipitation reached a maximum at different periods in different regions of China, with the trend of frontal migration paralleling the trend of summer insolation. However, this time-transgressive Holocene optimum was only supported by some coarse resolution records, especially from south China. He et al. (2004) reviewed multi-proxy paleoclimatic records across china and found that the Holocene optimum in eastern China was at ca. 6.5– 5.5 ka. Herzschuh (2006) assessed the late-Quaternary moisture change in Central Asia (including the Tibetan Plateau, northwest and northcentral China and Mongolia) using recent pollen data and other independent proxies. Her review suggested that the Holocene climate optimum with high precipitation occurred during the early Holocene in the Indian monsoon region, but possibly occurred during the midHolocene in the East Asian region. Nonetheless, the high-resolution speleothem isotope records with reliable chronology shed light on the general pattern of climate change during the Holocene in the monsoonal China region. However, the major patterns of temporal and spatial variation in past vegetation are far less clear than climate in this region. Pollen data would provide biological responses as well as climate change, in a widespread network compared to cave records, and would also help further test the Holocene pattern by using a unified synthesis methodology. Fossil pollen data from lakes and wetland deposits have been frequently used in vegetation and climate reconstructions, as they tend to reflect vegetation and climate changes at a regional scale. As a result, fossil pollen has become one of the most widely used and available paleoclimatic proxies. Liu (1988) reviewed late-Quaternary pollen data from temperate forests of China and found that the Holocene pollen stratigraphies suggest a tripartite division, with a period of maximum warmth, the Hypsithermal, in the mid-Holocene. Sun and Chen (1991) briefly reviewed the Holocene pollen data in China and indicated that vegetation change was sudden and remarkable in the early Holocene and late Holocene. Ren and Zhang (1998) and Ren and Beug (2002) presented Holocene pollen maps of China north of Yangtze River for six time slices at 2000-year intervals, revealing large changes in Holocene vegetation, especially since 6–4 ka. Yu et al. (1998) used a biomization approach to map pollen data and their reconstructions showed the shifts in biome distributions, implying significant changes in climate since 6000 yr ago. These syntheses are based on a large collection of Holocene pollen records of various time spans and data quality available in China; however, the chronologies are limited by dating material (most on bulk organic matter) and coarse dating control, and data sensitivity is also limited by the coarse sampling intervals and by the discontinuous nature of the sediment, including fluvial, coastal, or marine sediments, 243 255 255 used for pollen analysis (Liu, 1988). In addition, in the past couple of decades after these syntheses, many more high-resolution pollen records with relatively good chronology control have been available. Herzschuh (2006) assessed the late-Quaternary moisture change in Central Asia based on recent pollen data and other independent proxies, but her review evaluates very few records from eastern China. Hope et al. (2004) reviewed over 1000 marine and terrestrial pollen records to reconstruct the vegetation history in the Austrial-Asian region, including the monsoonal China; however, their analysis was mostly to examine vegetation change at broad regional and coarse temporal scales. There are no syntheses of pollen data specifically in the entire eastern monsoonal China for understanding the vegetation pattern and response to climate change. Therefore, there is a need to synthesize paleoclimatic information about monsoonal China based on recent published fossil pollen records. In this review, we used 31 fossil pollen records from eastern monsoonal China (see location in Fig. 1 and site information in Table 1). Our review and synthesis differ from earlier published reviews (e.g., Liu, 1988; Sun and Chen, 1991; Yu et al., 1998; An et al., 2000; Ren and Beug, 2002; Herzschuh, 2006), as we focused on monsoonal China, applied a uniform set of site-selection criteria, and used mostly recent pollen data. The objectives of this paper were to document regional vegetation patterns by synthesizing fossil pollen records from monsoonal China, and to evaluate and understand vegetation response to climate changes and the large-scale controls. This synthesis would not only bring together existing high-resolution data but also would assist the design of paleoecological and paleoenvironmental studies in the reviewed region. 2. Data sources and methods 2.1. Study region and site selection The geographical region considered in this review covers eastern monsoonal China (Fig. 1). Altitude ranges from near sea level to about 5000 m above sea level (Table 1). The Asian summer monsoon plays a significant role in controlling the effective moisture of the region. Under the influence of the monsoon systems, annual precipitation decreases sharply from the southeast to the northwest from >1800 mm to ca. 400 mm. At most of the sites the precipitation of the three summer months (JJA) (June, July, and August) usually accounts for more than 80% of the total annual precipitation, leaving winter and spring dry (Ren and Beug, 2002). The study sites are located in six vegetation zones (see details in Table 2; Fig. 2; Wu, 1980; Hou, 2001): 1) tropical monsoonal rain forest; 2) subtropical evergreen and deciduous forest; 3) temperate deciduous forest; 4) temperate conifer–deciduous mixed forest; 5) temperate steppe; and 6) highland meadow and steppe. Abundant pollen records of various time spans and data quality are now available in this region. However, many records tend to have low temporal resolution and can be discontinuous. In this study, we selected pollen sites based on three criteria: (1) a reliable chronology with a minimum of 4 dating control points over the Holocene; (2) high sampling resolution with a minimum 200 yr per sample; and (3) continuous record covering most of the Holocene without documented depositional hiatus. A total of 26 lake and peat sites met these criteria in this study. There are no satisfactory records in the temperate deciduous forest region. Therefore, we included two records with lower resolution in this region (the Yellow River Delta and Maohebei) when we discuss the general vegetation and climate history. In addition, three other sites (Poyang Lake, Taishizhuang and Tianchi Lake) that have high resolution pollen records but only for the 244 Y. Zhao et al. / Earth-Science Reviews 97 (2009) 242–256 Fig. 1. Map showing the location of fossil pollen sites in monsoonal China reviewed in this paper (see Table 1 for site information and references). late Holocene were also used to evaluate possible human disturbance on vegetation change. These three sites were not used in climate synthesis analysis owing to their limited time span. Our site-selection criteria make our review and synthesis different from earlier published reviews of pollen records (e.g., Liu, 1988; Sun and Chen, 1991; Ren and Zhang, 1998; Ren and Beug, 2002). Moreover, our synthesis includes the latest research results since 2000, covering more than 60% of the 31 pollen records used. Radiocarbon dating was the geochronological technique for all profiles used in this synthesis. All ages for the reviewed profiles have been calibrated or recalibrated to calendar years before the present (BP = AD 1950) using the latest IntCa04 calibration dataset (Reimer et al., 2004). Calibrated ages are used in compiling effective moisture curves and throughout the text (expressed as cal yr BP, or ka; 1 ka = 1000 cal yr BP). For the convenience of discussion, we organized the sites by six modern vegetation zones as described above. We present and discuss eight summary pollen diagrams that are representative of vegetation changes in these regions of different vegetation types. Pollen data were obtained from the original authors or digitized from pollen diagrams in the publications. At some sites, we also briefly describe the modern pollen-rain results, if available, to help evaluate fossil pollen interpretations. 2.2. Moisture index and statistical analysis The main pollen indices we used here include tropical tree pollen percentages, evergreen tree pollen percentages, broad leaved tree pollen percentages, and tree pollen percentage. We used these pollen indices to make a semi-quantitative estimate of relative moisture conditions, which is controlled by both precipitation and temperature (through influence on evaporation). The monsoonal region of China, including tropical monsoonal rain forest, subtropical evergreen forest and deciduous forest regions, often has high relative humidity in summer monsoon season, so evaporation would not increase significantly with temperature increase. Even in the steppe region near the limit of monsoon influence where evaporation might be an important factor for effective moisture, vegetation is more sensitive to minor change in effective moisture than the eastern parts of our study region. Therefore the monsoon-induced precipitation variability is more important than evaporation variability in understanding the variation in effective moisture in the monsoon region of China, as also indicated by An et al. (2000). As shown in other geological and biological records, effective moisture has a strong influence on vegetation in east-central China (An et al., 2000). We therefore used these pollen indices from 26 sites (excluding the two sites from the temperate deciduous forest region and three late Holocene sites) to estimate dry–wet climate fluctuations semi-quantitatively at individual sites during the Holocene. The four moisture classes were assigned from the wettest (score 4) to the driest period (score 1) at each individual site. The justification for designating an ordinal wetness scale is that pollen index types and values at each individual site may not be directly comparable to the ones at other sites, owing to different geographic locations, different dominant vegetation and hydrological settings. So the wetness scales are relative, in a semiquantitative sense, on the basis of individual sites. Here we use Qinghai Lake as an example to illustrate how we assigned relative wetness values to pollen records. We use total tree pollen (%) at Qinghai Lake as the index to infer effective moisture, with high tree pollen representing high tree dominance under a wet climate (Shen et al., 2005). Fossil pollen assemblages with total tree pollen percentages of <10%, 10– 20%, 20–30%, and 30–40%, receive wetness scores of 1, 2, 3 and 4, respectively. Although the score assignment is semi-subjective, it satisfactorily shows the general moisture trend at individual sites. Synthesized wetness curves for each individual region (vegetation zone) and the entire study region of monsoonal China during the Holocene were generated by averaging the wetness scores at 200-year time intervals for all 26 sites, with error bars as standard errors. Due to potentially different sensitivity to climate change at individual sites, the range and magnitude of percentage pollen change at individual sites can be very large (e.g., from <10 to >90% for total tree pollen), so averaging these highly variable values may not be reasonable and acceptable. Ordinal indices could smooth the range of variations for each site, thus providing more reasonable averaged values than original pollen data. This averaging procedure based on wetness scales assumes that each site is equally sensitive to climate change and equally well dated, and that there are no gradients in response across the region. In reality, some records were obviously better dated, at higher resolution, or higher sensitivity to climate change than others. Unfortunately, with information available in the original publications, it is not practical for the Y. Zhao et al. / Earth-Science Reviews 97 (2009) 242–256 245 Table 1 List of fossil pollen sites from eastern Asian monsoon China used in this review. Site no. Site name A1 Shuangchi Maar Lake E110°11′, N19°57′ Huguang Maar Lake E110°17′, N21°9′ Dahu E115°, N24°41′ Bajiaotian E110°20′, N25°48′ Chao Lake E117°16′54″, N31°25′28″ Poyang Lake E116°15′, N28°52′ Longquan Lake E112°1′, N30°52′ Dajiuhu E109°59′45″, N31°29′27″ Dianchi E 103°, N25° Erhai Lake E 100°05′, N25°36′ Shayema Lake E101°35′, N28°05′ Yellow River Delta E118°54.3′, N37°47.8′2 Maohebei E119°12′, N39°32′ Taishizhuang E115°49.5′, N40°21.5′ Jinchuan E126°22′, N42°20′ Gushantun E126°10′, N42°30′ Qindeli E133°15′, N48°00′ Haoluku E116°45.42′, N42°57.38′ Bayanchagan E115.21°, N41.65° Diaojiao Lake E112°21′ N41°18′ Daihai Lake E112°33′, N40°29′ Chasuqi E111°08′, N40°40′ Midiwan E108°37′, N37°39′ Tianchi Lake E106°19′, N35°16′ Dadiwan E105°54′, N35°01′ Zoige Basin E103°25′, N32°20′ Qinghai lake E99°36′, N36°32′ Hidden Lake E92°48′, N29°49′ Qongjiamong Co E92°22.37′, N29°48.77′ Zigetang Lake E90.9°′, N32.0° Selin Co E88°31′, N31°34′ A2 B1 B2 B3 B4 B5 B6 B7 B8 B9 C1 C2 C3 D1 D2 D3 E1 E2 E3 E4 E5 E6 E7 E8 F1 F2 F3 F4 F5 F6 Location Elevation Precip. Sample resolution Dating material (m a.s.l.) (mm) (yr) 75 1670 < 100 255 1600 250 Number of Type of date archives Reference Bulk organic matter 6 Lake core Zheng et al. (2003) 170 Bulk organic matter/leaves 7 Lake core Wang et al. (2007) 1600 240 Plant macrofossils 6 Peat section 1900 1814 250 Bulk organic matter 6 Peat core Xiao et al. (2007); Zhou et al. (2004) Li et al. (1993) 20 1000 < 200 14.1 1528 170 100 1100 < 150 1760 1500 1886 Bulk organic matter /charcoal/ 10 shell/ plant macrofossil Bulk organic matter 9 Fuvial + lake sediment Lake core Chen et al. (2009) Bulk organic matter 6 Lake core Liu et al. (1993) 120 Peat cellulose 7 Peat section Zhu et al. (2006) 1010 160 Bulk organic matter 7 Lake core Sun et al. (1986) 1974 900 < 100 Bulk organic matter 6 Lake core Shen et al. (2006) 2400 1090 130 Bulk organic matter 5 Lake core Jarvis (1999) 4.9 750 <1000 Mollusc shell 9 Delta borehole Yi et al. (2003) 2 650 > 300 Bulk organic matter 4 Peat section Li and Liang (1985) 500 420 < 100 Bulk organic matter 6 Peat section Tarasov et al. (2006) 662 700 210 Bulk organic matter 4 Peat section Jiang et al. (2008) 600 630 300 6 Peat section Liu (1989) 52 600 280 Bulk organic matter, charcoal Bulk organic matter 5 Peat section Xia (1988) 1295 370 200 Bulk organic matter 4 Lake section Liu et al. (2002) 1355 400 130 Bulk organic matter 7 Lake core Jiang et al. (2006) 1800 421 85 Bulk organic matter 4 Lake core 1221 423 < 100 Bulk organic matter 8 Lake core Shi and Song (2003); Song et al. (1996) Xiao et al. (2004) 1000 400 70 Bulk organic matter 4 Peat section Wang and Sun (1997) 1400 395 100 Bulk organic matter 23 Peat section Li et al. (2003) 2430 615 85 Plant macrofossil 19 Lake core Y. Zhao (unpublished data) 1400 400 <50 Bulk organic matter 3 Marsh section An et al. (2003) 3492 710 190 Bulk organic matter 8 Peat section Yan et al. (1999) 3200 350 60 Bulk organic matter 7 Lake core Shen et al. (2005) 4980 450 110 Plant macrofossils 7 Lake core Tang et al. (2000) 4980 450 < 200 20 Lake core Shen (2003) 4560 320 160 Bulk organic matter 5 Lake core Herzschuh et al. (2006) 4530 290 200 Bulk organic matter 5 Lake core Sun et al. (1993) purpose of this synthesis to assign weightings to each individual site. Nonetheless, these synthesized time series help us document and understand variations of effective moisture, though with some uncertainties as shown in the error bars for the synthesis curve. We choose the squared-chord distance (SCD), which has proven especially satisfactory to reveal regionally synchronous palynological changes from pollen-frequency data (e.g. Overpeck et al., 1985; Grimm and Jacobson, 1992; Faison et al., 2006). SCD is calculated as: Flies/charcoal/macrofossil Jiang and Piperno (1999) 1/2 2 dij = σk(p1/2 ik − pjk ) , where dij is the SCD between two multivariate samples i and j, and pik is the proportion of species k in sample i. For each pollen time series (site) used in this synthesis, the pollen data were interpolated at 200-year intervals by using the program AnalySeries 2.0.4 (Pailard et al., 1996). The SCD between the adjacent 200-year samples was used to calculate the rate of palynological change per 100 yr (Grimm and Jacobson, 1992). Although most of the selected pollen records have only 5–10 radiocarbon dates, the age 246 Y. Zhao et al. / Earth-Science Reviews 97 (2009) 242–256 Table 2 Modern environmental settings (Wu and Zhang, 1992). Vegetation type MAT (°C) MAP (mm) Dominant plant taxa Tropical monsoonal rain forest Subtropical evergreen and deciduous forest Temperate deciduous forest Temperate conifer– deciduous mixed forest Temperate steppe 22 to 26.5 14 to 22 >1000 9 to 14 500–900 2 to 8 500–800 − 3 to 8 200–450 − 2 to 0 300–400 Terminalia hainanensis, Lannea coromandelica, Ficus altissima, Chukrasia tabularis, Vatica astrotricha, Madhuca subquincuncialis, Altingia chinenesis, Dacrydium pierrei, Lithocarpus fenzelianus, and Adinandra hananensis. Quercus variabilis, Q. serrata, Q. fabri, Castanopsis sclerophylla, Castanea sequinii, Platycarya strobilacea, Lithocarpus cleistocarpus, Betula luminifera, and Liquidambar formosana. Quercus mongolica, Q. liaotungensis, Q. aliena, Q. acutissina, Q. variabilis, Robinia pseudoacacia, Salix matsuduna, and Populus simorill. Pinus koraiensis, P. sylvestris, Abies nephrolepis, Picea jezoensis, Larix gmelinii, L. olgensis and Quercus mongolica, Tilia amurensis, Ulmus propinqua, and Betula platyphylla. Artemisia spp., Stipa spp., Carex spp., Rosaceae and Fabaceae. Pinus, Quercus, Betula, Ulmus, Salix, Corylus and Populus are the main tree taxa on nearby hills. Highland meadow: Cyperaceae (e.g. Kobresia pygmaea), Poaceae (such as Achnatherum splendens, Stipa breviflora, S. gobica). Asteraceae, Rhododendron, Gentianaceae, Fabaceae, Saxifragaceae and Polygonum; highland steppe: more Poaceae (e.g. Achnatherum splendens, Stipa breviflora, S. gobica) and Artemisia frigida, A. scoparia than alpine meadow. Highland meadow and steppe 1200–3000 MAT = mean annual temperature; MAP = mean annual precipitation. controls should be sufficient to detect abrupt changes at multicentennial or millennial scales. Our site-selection criterion of continuous record covering most of the Holocene without documented depositional hiatus also makes the analysis justifiable. The analysis does not reveal the qualitative nature of climate change, but it identifies times of rapid change regardless of the direction. Moreover, the average for a large region will not identify timetransgressive change, but it reveals abrupt, synchronous, geographically widespread change (Huntley, 1992). 3. Pollen records of Holocene vegetation and climate changes Below we will describe the synthesis results of vegetation and climate change from each of six main vegetation regions in monsoonal China. 3.1. Tropical monsoonal rain forest Two high-resolution pollen records are available from the tropical and southern subtropical regions of southeast China. One record is from Shuangchi Maar Lake (site A1) on Hainan Island, spanning the last ca. 9000 yr (Zheng et al., 2003). The pollen assemblages were dominated by tropical trees, mainly including Mallotus and Casearia at 8.9–4.3 ka, suggesting a warm and wet climate. After 4.3 ka, herb and fern pollen increased at the expense of tree pollen, and total pollen concentration was very low, indicating a drying climate. The other pollen diagram from Huguangyan Maar (site A2) has a sampling resolution of ca. 170 yr and is divided into 3 zones (Fig. 3A; Wang et al., 2007). The percentage of tropical tree pollen is high during 11.6–7.8 ka, mainly including Moraceae, Mallotus, Ficus and Aporosa. The tropical vegetation represented by pollen suggests a warm and wet climate. From 7.8 ka, tropical tree pollen started to decrease, especially after 4.2 ka. 3.2. Subtropical evergreen and deciduous forest More well dated high-resolution pollen records are available from this region. A peat profile at Dahu peatland (site B1) provided a well dated Holocene pollen record (Fig. 3B; Zhou et al., 2004; Xiao et al., 2007). The pollen diagram indicates a major change dominance of Alnus to evergreen tree pollen of Castanopsis/Lithocarpus at 10.4 ka, signifying a warming and wetting climate. Castanopsis/Lithocarpus is sclerophyllous taxa that are adapted to spring drought, compared to Fig. 2. Dominant biomes (vegetation types) in China (after Wu, 1980). Y. Zhao et al. / Earth-Science Reviews 97 (2009) 242–256 Fig. 3. Summary percentage pollen diagrams from eastern monsoonal China. (A) Huguangyan Maar Lake (A2), South China (redrawn from Wang et al. (2007)); (B) Dahu peatland (B1), South China (redrawn from Xiao et al. (2007)); (C) Dajiuhu peatland (B6), Central China (redrawn from Zhu et al. (2006)); (D) Shayema Lake (B9), southwestern China (redrawn from Jarvis (1993); (E) Maohebei peatland (C2), north China (redrawn from Li and Liang (1985)); (F) Qindeli peatland, northeastern China (redrawn from Xia (1988)); (G) Bayanchagan Lake (E2), northwestern China (redrawn from Jiang et al. (2006)); (H) Qinghai Lake (F2), northeastern Tibetan Plateau (redrawn from Shen et al. (2005)). All the percentages were calculated based on terrestrial pollen sums. 247 248 Y. Zhao et al. / Earth-Science Reviews 97 (2009) 242–256 many mesic deciduous forest taxa; however, we interpret its presence as indicating warm/wet condition as it requires more moisture than Alnus and herbs that are abundant in these pollen records. At 10.4– 6 ka, Castanopsis/Lithocarpus remains high, reaching a maximum of near 30%. Cyclobalanopsis pollen increases gradually to a value of 19% during that period. This period marks the Holocene optimum period (warmer and wetter climate), as represented by a subtropical evergreen broadleaved forest. After 6 ka, the pollen record shows a rapid increase in ferns and herbs at the expense of tree pollen, suggesting a drier climate. Pollen record from Caohu Lake (B3), west to the Yangtze River Delta, indicated an evergreen and deciduous mixed broadleaved forest dominated by Cyclobalanopsis and Quercus after 10.5 ka, especially between 8.2 and 7.5 ka, suggesting a warm and wet climate. At 7.5– 3.7 ka, there are noticeable fluctuations in main pollen taxa, in particular a general decline in Cyclobalanopsis and other arboreal pollen and an increase in terrestrial herbs. After 3.7 ka, the broadleaved forest trees largely gave way to terrestrial herbs. Pinus continued to increase alongside with most herbs until 2 ka (Chen et al., 2009). Pollen record at ca. 120-year sampling resolution from the Dajiuhu peatland (site B6), in Shennongjia Mountains in central China (middle reach of Yangtze River), shows a progressive increase in percentages of evergreen tree pollen after 11 ka (Fig. 3C; Zhu et al., 2006). During 11– 6 ka, pollen assemblages are marked by a continuous increase in evergreen components as well as some deciduous tree pollen, such as Cyclobalanopsis/Castanopsis, Betula, Carpinus, Quercus and Juglans. Modern pollen assemblages from surface samples in the Shennongjia region are dominated by deciduous broadleaved trees and represent regional pollen source (Liu et al., 1993). There are good relationships between pollen and climate variables, with higher evergreen tree pollen percentages at warm and wet sites (Liu et al., 2000; Zhu et al., 2008). The fossil pollen assemblages imply a mixed forest of deciduous and evergreen broadleaved trees under a warming and wetting climate during the early and mid-Holocene. After 4.5 ka, coniferous tree pollen increased at the expense of evergreen tree pollen, suggesting mixed forests of coniferous trees with deciduous broadleaved trees caused by a weakening of the East Asian summer monsoon. Another pollen record from Longquan Lake in central China (site B5; Li et al., 1993) similarly revealed that evergreen broadleaved — deciduous broadleaved forest (Castanea, Hamamelis, Keterleeria, and Tsuga) dominated from 10.7 to 4.6 ka, while herb became dominant at the expense of tree pollen after 4.6 ka. The pollen record from Shayema Lake (site B9) in southwestern China spans the last 13,000 yr (Fig. 3D; Jarvis, 1993). Modern pollen-rain samples were collected from surface sediments at 13 lakes along an elevational transect between 1800 and 3900 m above sea level (Jarvis and Clay-Poole, 1992). The results show that a significant number of important taxa in the region are insect-pollinated and produce minimal pollen grains. To mitigate the difficulty of working with such a large number of underrepresented taxa, pollen percentage values for low pollen-producing taxa of similar autoecological features (e.g., mesic deciduous, sclerophyllous evergreen broadleaved) were summed and examined together, by giving particular weight to the presence of key pollen types representative of a particular climatic regime. The fossil pollen record shows several changes in the vegetation and climate since 13 ka (Jarvis, 1993). From 12.7 to 10.6 ka, cold-tolerant species, such as Abies, Betula, and deciduous oaks (Quercus), dominated the vegetation. Between 10.6 and 7.9 ka, the abundance of deciduous oaks decreased, while evergreen oaks increased, as did Tsuga and mesic deciduous species, suggesting a warm climate with probably increased precipitation. During the period 7.9–4.7 ka, spring drought-adapted sclerophyllous taxa (such as Lithocarpus/Castanopsis) increased at the expense of mesic deciduous taxa, indicating that precipitation was becoming more seasonal. This is in a great contrast to the climate in the early Holocene, with a reduced seasonality in precipitation and an increased seasonality in temperature, when winters were colder and spring moisture was probably more abundant. The drought-adapted tree taxa continued to be the dominant components of the vegetation until ca. 1 ka when the effects of human disturbance are noted in the pollen record (Jarvis, 1993). Evidence for deforestation is supported by an increase in Alnus, Artemisia, Poaceae, Pteridium, and Pteris and the minimal amounts of Tsuga and Picea. Other palynological records from southwestern China (Dianchi: site B7, Sun et al., 1986; Erhai: site B8, Shen et al., 2006) also suggest that prior to 10 ka the climate was colder and drier relative to today as inferred from dominant conifer pollen (Pinus, Abies). From the early to middle Holocene, pollen assemblages show a gradual increase in abundance and diversity of mesic deciduous and evergreen trees (Quercus, Tsuga, Castanopsis/Lithocarpus). Vegetation was reflected by spring drought-adapted sclerophyllous taxa relative to mesic deciduous taxa during the late mid- to late Holocene. At Erhai Lake, Pinus increased significantly, along with pollen of Plantago, Artemisia, Chenopodiaceae and Poaceae, during late Holocene, suggesting human disturbance (Shen et al., 2006). 3.3. Temperate deciduous forest There are few pollen records with good chronology and high resolution from this region. Palynological analyses of two boreholes from the Yellow River Delta (site C1) provide a vegetation and climate history of the last 12 ka, but only on millennial scale (Yi et al., 2003). From 9.8 to 4.5 ka evergreen and broadleaved deciduous forest thrive (indicated by high Quercus, Carpinus/Ostrya, Ulmus/Zelkova pollen values), suggesting warmer and wetter climatic conditions. A significant reduction in deciduous Quercus pollen and an increase in conifer Pinus pollen at ca. 4 ka, together with the first appearance of buckwheat (Fagopyrum) pollen at 1.3 ka, probably reflect widespread human disturbance of the natural vegetation and intensive cultivation. Another record with >300-year sampling resolution from Maohebei peatland (site C2) spanned from 12.5 to 3 ka (Fig. 3E; Li and Liang, 1985). The pollen record shows that deciduous tree pollen (represented by Quercus, Tilia, Ulmus, Carpinus, Corylus and Betula) peaks at ca. 11.5–8.5 ka, corresponding to warm and humid periods, and the subsequent interval of lower percentages of these trees, corresponding to a dry climate interval (Li and Liang, 1985). 3.4. Temperate mixed conifer–hardwood forest Three pollen records from peat profiles (Jinchuan: site D1, Jiang et al., 2008; Qindeli Bog: site D3, Xia, 1988; Gushantun Bog: site D2, Liu, 1989) in the mixed conifer–hardwood forest of northeast China span the entire Holocene. Pollen record at ca. 280-year resolution from Qindeli shows that from 12.5 to 10 ka broadleaved trees, including Betula, dominated the vegetation (Fig. 3F). At 10–5.5 ka, deciduous broadleaved forest trees, including Ulmus, Quercus, Salix, Carpinus, and Corylus, thrived, indicating a high effective moisture during the growing season. After 5.5 ka, Pinus and Abies/Picea increased at the expense of deciduous broadleaved tree pollen, suggesting a conifer-dominated forest and thus a colder and drier climate. This trend was accentuated after 2.5 or 2 ka as indicated by further expansion of the boreal conifers. At Gushantun Bog, Betula increased at ~11.3 ka, and from 11.3 to 4.4 ka, broadleaved tree pollen from Quercus, Ulmus, Alnus, Juglans and Corylus had high values. Since 4.4 ka, Pinus increased while deciduous broadleaved trees decreased, especially after 2.2 ka. The Jinchuan profile has a similar pattern as Qindeli, with a steadily high broadleaved pollen (Quercus, Ulmus and Juglans) from 11.5 to 6/5 ka, suggesting an interval of high effective moisture. 3.5. Temperate steppe Jiang et al. (2006) presented a pollen record at ca. 130-year sampling resolution from Bayanchagan Lake in Inner Mongolia (site E2; Fig. 3G). The pollen assemblages show that vegetation around Bayanchagan Lake changed from a steppe at 12.5–9.2 ka, through a Betula/Pinus-dominated Y. Zhao et al. / Earth-Science Reviews 97 (2009) 242–256 steppe woodland at 9.2–6.7 ka, back to steppe after 6.7 ka. Jiang et al. (2006) used standard modern analogue technique to reconstruct changes in paleoclimatic parameters based on modern pollen datasets (including 211 surface pollen assemblages from northern China) and corresponding climate data. The vegetation sequence and paleoclimatic reconstruction suggest that a relatively humid climate during the early mid-Holocene at 9.2–6.7 ka was favorable for the development of woodland. Pollen assemblages at other sites (Haoluku: site E1, Liu et al., 2002; Daihai Lake: site E4, Xiao et al., 2004; Diaojiao Lake: site E3, Shi and Song, 2003; Chasuqi: site E5, Wang and Sun, 1997) in eastern Inner Mongolia are generally dominated by Betula, Picea, and Ulmus during the early Holocene, while by Artemisia with some tree pollen including Betula, Quercus, Ulmus, and Pinus in the middle Holocene from 7 to 5 ka. After 5 ka or 4 ka, pollen assemblages were dominated by Artemisia with some tree pollen. Vegetation changed from steppe at the beginning of Holocene, to forest steppe in the late part of early Holocene and mid-Holocene, to steppe in the late Holocene. These vegetation sequences suggest that climate changed generally from a wet climate in the early and mid-Holocene to drier conditions after ca. 5 ka. Dadiwan marsh section (site E8), located in the NW Loess Plateau, has a high sampling pollen resolution of ca. 50 yr (An et al., 2003). The pollen assemblages show that vegetation around Dadiwan changed from a desert steppe at 10–8.5 ka, through a Pinus-dominated steppe woodland at 8.5–6.4 ka, to back to desert steppe after 6.4 ka. The vegetation sequence suggests that a relatively humid climate during the mid-Holocene at 8.5–6.5 ka was favorable for the development of woodland. Midiwan (site E6; Li et al., 2003), another site on the NW Loess Plateau, shows a different pattern of Holocene vegetation and climate change. Vegetation was sparsely wooded grassland, consisting mainly of Thalictrum, Cyperaceae, Betula, and Quercus at 11.5–8.5 ka, indicating a warm and humid early Holocene. 3.6. Highland meadow and steppe Six pollen diagrams from this region (from sites F1 to F6) were used for this synthesis. The pollen diagram from Qinghai Lake (site F2; Fig. 3H; Shen et al., 2005) can be divided into three pollen assemblage zones. Before 10.6 ka, pollen assemblages were characterized by Artemisia (~60%), tree pollen (~20%), Poaceae (~10%), and Chenopodiacece (~10%). During 10.6 and 4.2 ka, Artemisia decreased to <40%, Pinus increased up to ~30% and Betula was up to 20%. The highest pollen concentrations occurred in this zone. After 4.2 ka, Artemisia increased again (up to 80%) while Pinus decreased to 15–0% (Shen et al., 2005). Vegetation around Qinghai Lake changed from steppe before the Holocene, through steppe forest in the early and mid-Holocene at 10.6–4.2 ka, to Artemisia-dominated steppe in the late Holocene. This vegetation sequence suggests a dry climate before the Holocene, a wet climate in the early and mid-Holocene, and a dry climate in the late Holocene. Pollen assemblages from other sites in the Tibetan Plateau (Zoige Basin: F1, Yan et al., 1999; Hidden Lake: F3, Tang et al., 2000; Co Qongjiamong: F4, Shen, 2003; and Selin Co: F6, Sun et al., 1993) were dominated by Artemisia and Cyperaceae, with relatively high Pinus/Picea and Betula pollen percentages during the early and middle Holocene from 11.5 to 6 or 5 ka, while dominated by Artemisia, Cyperaceae and Poaceae during the late Holocene. The vegetation sequence suggests a humid early and mid-Holocene and a dry late Holocene. However, at Zigetang Lake (F5), pollen assemblages are consistently dominated by Artemisia and Cyperaceae (Herzschuh et al., 2006). Pollen assemblages suggest a dominance of temperate steppe vegetation during the first half of the Holocene, while alpine steppes with desert elements tend to dominate the second half. The vegetation sequence indicates a general cooling and drying trend throughout the Holocene. 249 4. Discussion 4.1. Temporal and spatial patterns of Holocene vegetation and climate change The major vegetation changes at individual sites from various vegetation zones show different types and patterns during the Holocene (Fig. 4). In the tropical monsoonal rain forest region, vegetation changed from seasonal rain forest dominated by tropical trees (e.g., Fig. 5A) to more open forest. In the subtropical region, vegetation changed from evergreen broadleaved forest dominated by evergreen trees Quercus and Castanopsis/Lithocarpus (e.g., Fig. 5B, C and D), to evergreendeciduous broadleaved forest dominated by Quercus, Betula, Carya, and Ulmus or mixed forest dominated by Pinus and deciduous trees with more herbs during the late Holocene. In the temperate deciduous forest region, vegetation shifted from deciduous broadleaved forest during early and mid-Holocene (e.g., Fig. 5E), to diverse broadleaved deciduous forest with some herbs (Artemisia, Poaceae) during the late Holocene. In the conifer–hardwood mixed forest region, vegetation changed from deciduous broadleaved forest (Quercus-dominated) to conifer–deciduous mixed forest dominated by conifer and other trees (Pinus, Picea, Abies, Quercus and Betula) (e.g., Fig. 5F). In the temperate steppe region, steppe (Artemisia, Poaceae), forest steppe (Betula, Pinus, and Artemisia), and steppe dominated the vegetation over the Holocene in eastern Inner Mongolia (e.g., Fig. 5G). Vegetation in the northwestern Loess Plateau changed between desert steppe (Artemisia, Chenopodiaceae, and Poaceae), forest steppe (Pinus, Artemisia, Poaceae, and Asteraceae) and steppe. In the highland meadow and steppe region, vegetation is characterized by meadow/steppe (Artemisia, Poaceae, and Cyperaceae) and steppe forest (mainly Pinus, Picea and Abies) in the early and midHolocene (e.g., Fig. 5H) and meadow/steppe during the late Holocene. The semi-quantitative moisture classes obtained from interpreted fossil pollen records in the six regions reveal generally synchronous wet–dry climate changes during the Holocene (Fig. 6), though their vegetation shows different types and patterns. A wet climate occurred in the early and mid-Holocene before 6–5 ka, with the maximum moist period at 9.5–6 ka. All the records show a consistently dry climate during the late Holocene, with an abrupt shift at ca. 4.5 ka (Fig. 6G). Due to the relative lack of fossil pollen records in the regions of tropical monsoonal rainforest and temperate deciduous forest, the pattern in these regions at the beginning of Holocene is not as clear as in other regions. Vegetation change at individual sites (Fig. 4) and the moderate error of the moisture curve (Fig. 8E) together indicate that the analysis results of generally synchronous patterns in different regions are not an artifact that could be caused by averaging approach. An et al. (2000) proposed that the Holocene optimum, as defined by maximum East Asian summer monsoon precipitation, was asynchronous in different regions, at ca. 10–8 ka in northeastern China, at 10–7 ka in north-central and north east-central China, at ca. 7–5 ka in the middle and lower reaches of the Yangtze River, at ca. 3 ka in southern China, and at 11 ka in southwestern China. However, our synthesis based on pollen data and other evidence from recently published records does not appear to support this hypothesis (also see Feng et al., 2006). The different conclusions from these syntheses are partly due to the fact that An et al. (2000) used some records with low sampling resolution and poor dating controls, especially those from southern China. Dissimilarity analysis of pollen data using SCD measures for the entire eastern monsoonal China shows that times of significant palynological change occurred at 11–10 ka, 6–5 ka and 2–1 ka (Fig. 7). The regional average curve, which smoothes out site-specific changes, can reveal times of regionally synchronous palynological changes in a large geographic region. Our result generally agrees with that inferred from T-test analysis based on multi-proxy records in Asian monsoon region (Morrill et al., 2003). At the start of the Holocene, the average rate of change is high at most sites due to the transition from the late-glacial period to the Holocene. The significant changes during this time result 250 Y. Zhao et al. / Earth-Science Reviews 97 (2009) 242–256 Fig. 4. Summary of vegetation history derived from fossil pollen records in eastern monsoonal China. See Table 1 for site information and references. Y. Zhao et al. / Earth-Science Reviews 97 (2009) 242–256 Fig. 5. Summary of major vegetation taxa change in various regions arranged from south to north in eastern monsoonal China. (A) Tropical tree pollen (%) at Huguang Maar Lake (Wang et al., 2007); (B) evergreen tree pollen (%) at Dahu (Zhou et al., 2004); (C) evergreen tree pollen (%) at Dajiuhu (Zhu et al. (2006)); (D) sclerophyllus tree pollen (%) at Shayema Lake (Jarvis, 1993); (E) deciduous tree pollen (%) at Maohebei peatland (Li and Liang, 1985); (F) conifer tree pollen (%) at Qindeli peatland (Xia, 1988); (G) tree pollen (%) at Bayanchagan Lake (Jiang et al., 2006); (H) tree pollen (%) at Qinghai Lake (Shen et al., 2005). 251 252 Y. Zhao et al. / Earth-Science Reviews 97 (2009) 242–256 Fig. 6. Synthesized time series of relative moisture changes as inferred from fossil pollen data across eastern monsoonal China. (A) Tropical monsoonal rain forest (n = 2); (B) subtropical evergreen forest (n = 8); (C) temperate forest (n = 2); (D) temperate mixed forest (n = 3); (E) temperate steppe (n = 7); (F) highland meadow and steppe (n = 6); (G) eastern monsoonal China (n = 26). The moisture conditions were averaged at every 100 yr and coded by four classes: 1 — dry; 2 — moderately dry; 3 — moderately wet; and 4 — wet. mostly from expansions of various tree populations in eastern monsoonal China. Between 10 and 8 ka, the vegetation became relatively stabilized. The climate change was probably too slow or too small (Fig. 8) to cause synchronous regional response to vegetation. Then major changes were initiated at many sites beginning at about 6–5 ka, owing mostly to tree decline (Fig. 4). Maximum rate of summer insolation change occurring during the mid-Holocene (Berger and Loutre, 1991) might have caused monsoon intensity shift (Wang et al., 2005) and then strong vegetation changes during that time (see discussion in Section 4.2). Another high peak of SCD occurred at around 2–1 ka, due to peak values of Pinus and herbs at many sites. Deforestation could have caused relatively large changes in vegetation over the last 2 ka (Fig. 9), as discussed in Section 4.3. 4.2. Vegetation responses to climate oscillations Various proxy data from speleothem, lake sediments and peat cores show similar patterns in monsoon intensity change during the Holocene in monsoonal China (Fig. 8; Hong et al., 2003; Wang et al., 2005, 2008; Hu et al., 2008). At Dongge Cave, oxygen isotope values show a major negative shift at the end of the Younger Dryas at 11.5 ka, reaching a minimum at ca. 9 ka, followed by a gradual long-term increase. Isotope has been used to track past summer monsoon intensity (Wang et al., 2005). Therefore the oxygen isotope at Dongge Cave shows a strong Asian monsoon period in the early Holocene, followed by a gradual weakening trend since 7 ka. Further north at Sanbao Cave and Heshang Cave, oxygen isotopes show the similar Fig. 7. Average rate of palynological change during the Holocene from all the fossil pollen records (n = 26) in eastern monsoonal China. Y. Zhao et al. / Earth-Science Reviews 97 (2009) 242–256 253 Fig. 8. Comparison of synthesized Holocene effective moisture evolution with other selected proxy records from eastern monsoonal China. (A) Oxygen isotope at Dongge Cave (Wang et al., 2005); (B) oxygen isotope at Sanbao Cave (Shao et al., 2006; Wang et al., 2008); (C) oxygen isotope from ostracode shells at Qinghai Lake (Liu et al., 2007). (D) carbon isotope from Hongyuan peat; (E) pollen-based moisture index in monsoonal China, with error bars as standard errors; (F) summer insolation at 30°N latitude (Berger and Loutre, 1991). general trend as Dongge Cave during the Holocene. Additional lines of evidence of isotope from lake sediment (at Qinghai Lake) (Liu et al., 2007) and peat (at Hongyuan and Hani sites) (Hong et al., 2003, 2005) and lake level records (Xue and Yu, 2000) support the conclusions drawn from the cave records, indicating a marked stronger summer monsoon phase in the first half of the Holocene and a gradual decrease during the second half, until the weakest summer monsoon during the late Holocene. The general trend of Holocene Asian monsoon history has been attributed to changes in summer insolation at low latitudes (Kutzbach, 1981; COHMAP, 1988; Wang et al., 2005). Strong summer insolation in the Northern Hemisphere during the early Holocene (Berger and Loutre, 1991) induced strong land–ocean pressure and temperature gradients and increased onshore moist air flow in the summer, causing an enhanced Asian summer monsoon (COHMAP, 1988). The gradual weakening of the Asian summer monsoon since the mid-Holocene was in response to the orbitallyinduced decrease in summer insolation (e.g., Gupta et al., 2003), enhanced by the feedbacks from changes in vegetation cover and soil moisture as was the case in North Africa (Kutzbach et al., 1996; Ganopolski et al., 1998). Fig. 9. Sites with evidence for human activity from fossil pollen data. Numbers indicate the ages of records (ka). 254 Y. Zhao et al. / Earth-Science Reviews 97 (2009) 242–256 The climate pattern inferred from fossil pollen records in eastern monsoonal China generally correlates with the monsoon intensity history as discussed above, namely moist phase in the first half of the Holocene and a gradual decrease in effective moisture during the second half (Fig. 8). At 9.5–6 ka, moisture based on pollen data in eastern monsoonal China reached the maximum. Therefore our synthesis demonstrates that changes in vegetation closely correlate with independently documented changes in climate revealed by other proxies. However, during the first two millennia in the Holocene from 11.5 to 9.5 ka, effective moisture inferred from fossil pollen records show only a slow and gradual increase, while most independent climate records, for example from Dongge Cave and Sanbao Cave, tend to show an abrupt onset of the maximum monsoon. Several reasons might be responsible for that apparently gradual response or 1000-year time lag in vegetation. (1) Vegetation, especially tree-dominated forests, tends to show 100–200 yr lag, as mature trees are less sensitive to climate than seedlings (Williams et al., 2002). However, it is difficult to explain the 1000-year lag, considering that this region was not glaciated, so vegetation immigration delay often-documented in North America and Europe is not applicable here (Davis, 1986; Lang, 1994; Williams et al., 2002). (2) A mismatch of favorable moisture and temperature conditions could cause that delay, if temperature was not favorable in the early Holocene (Shi et al., 1992), even though precipitation is high (Wang et al., 2005). (3) Age uncertainty of pollen records from lake and peat sediments could also cause that apparent time lag or gradual response. Effective moisture inferred from pollen data started to decrease from 6 ka, with an abrupt shift at ca. 4.5 ka. Rates of palynological change also demonstrate an abrupt change at around 6–5 ka. Climate change may have been responsible for this vegetation shift. Firstly, independent isotope records from Dongge Cave (Fig. 8A; Wang et al., 2005) and Sanbao Cave (Fig. 8B; Shao et al., 2006; Wang et al., 2008) show that since 6 ka or 5 ka monsoon intensity decreased, suggesting that vegetation change around that time was likely forced by natural factors. These cave records also show an abrupt lowering of Asian monsoon intensity at ca. 4.4 ka over several decades. In addition to the indication by other independent climatic proxies as discussed above, climate drying was also supported by the evidence of the cultural responses to prolonged drought during the late Holocene (particularly around 4.5–4 ka) in Asia and Africa, including population dislocations, urban abandonment and state collapse (deMenocal, 2001), for example, the collapse of Neolithic culture around the Central Plain in China (Wu and Liu, 2004). Secondly, pollen evidence of clear human disturbance is lacking before 2 ka, except at the sites from the Yangtze River drainage basin (Fig. 9). For example, at Taishizhuang (site C3), located in temperate deciduous forest region, both archaeological and paleoenvironmental data support the conclusion that changes in pollen composition between 5.7 and 2.1 ka reflect natural variation in precipitation but not deforestation caused by humans (Tarasov et al., 2006). Archaeological records from 100 sites show the habitation of northeastern China during the prehistorical and early historical periods since ca. 8.2 ka, but do not provide evidence for the use of wood resources or for widespread farmlands that would influence the regional vegetation development and leave traces in the pollen assemblages. 4.3. Human disturbance during the late Holocene Although we argue that insolation-induced change in monsoon intensity and precipitation was responsible for vegetation changes during the late Holocene, human disturbance also played a major role and contributed to the abrupt vegetation shift over the last several millennia, especially over the last 2000 yr. In China, there lacks a regionally distinct, broadly synchronous, and clearly defined pollen marker horizon for human disturbance (Liu and Qiu, 1994), or settlement comparable to the Ambrosia-rise in North America (McAndrews, 1988), or the Ulmus decline in western Europe (Behre, 1981). Fig. 9 indicated the timing of some sites in eastern monsoonal China that are exposed to human activities inferred from fossil pollen records. It shows that there is some pollen evidence indicating human disturbance on vegetation in the south part of monsoonal China. Atahan et al.'s (2008) work in the lower Yangtze River region showed that substantial human impacts are evident at 4.7 ka, 4.1 ka and 2.4 ka at three sites in that region, as suggested by wild rice (Oryza pollen and Oryza phytoliths). Detailed evidence of pollen, algal, fungal spore and micro-charcoal data from sediment at Kuahuqiao from the Yangtze Delta reveal the precise cultural and environmental context of rice cultivation at this earliest known Neolithic site in eastern China at 7.7 ka (Zhong et al., 2007). At Poyang Lake in the subtropical region, strong human influence over the last 2 ka was inferred from highresolution pollen record during the late Holocene, indicated by significant reduction of arboreal pollen and an increase of herbaceous pollen (Jiang and Piperno, 1999). Two high-resolution pollen diagrams (Shayema Lake and Dianchi Lake) from southwest China show that the impact of human settlement and agriculture was only clearly expressed in the sediment and pollen records after 1.5–1 ka by an increase in clastic contents and the occurrence of disturbance indicators (Phyllanthus, Plantago, Poaceae, Pteridium) and cultigens (Cannabis, Fagopyron). Another pollen record from Erhai Lake in southwest China shows a pronounced phase of deforestation since 2.2 ka, characterized by a rapid decline in Pinus and a marked increase in Poaceae along with other disturbance taxa, including Epilobium herbs (Shen et al., 2006). At Tianchi Lake (E7) from the southwestern Loess Plateau, pollen records show that vegetation changed from forest to steppe-like vegetation at ca. 2 ka and Poaceae pollen and microscopic charcoal both show sharp increase since then (Y. Zhao, unpublished data), suggesting that human activities over the last 2000 yr have significantly accelerated deforestation that was initiated by a drying climate since the mid-Holocene at Tianchi Lake. Most pollen records reviewed here are from lake sediments and reflect broad regional vegetation changes. Human activities might have affected vegetation at some local sites early in the Holocene, but there is no evidence showing consistent vegetation change at regional scales induced by human activities at that time. In any case, the fossil pollen records revealed synchronous deforestation most likely through slash-and-burn practice for agriculture development or wood resource at least around 2 ka (Jiang et al., 2008). In addition to agriculture and wood usage, grazing is another important human activity that affects vegetation in northern China. Some studies (e.g., Miehe et al., 2008; Schlütz and Lehmkuhl, 2009) on nomadic influence on vegetation have been undertaken on the Tibetan Plateau that was previously thought to be less influenced by human activities. Miehe et al. (2008) proposed that the presently degraded pastures of the northeastern Tibetan Plateau largely originate from forests since at least 8 ka based on pollen indicators. Schlütz and Lehmkuhl (2009) found that first signs of nomadic presence appear as early as 7.2 ka and the Poaceae-rich natural vegetation was transformed by nomadic grazing to Kobresia-pastures at 5.9–2.7 ka on the Nianbaoyeze Mountain in the eastern margin of the Tibetan Plateau, based on a suite of geomorphological and palynological data. However, these data from the northeastern and eastern Tibetan Plateau are from soil profiles without robust age controls and represent localized vegetation signals, which are not suitable for detecting large-scale vegetation change induced by human activity as from lake sediments. Nevertheless, much more research with focus on the grazing history is needed to establish a coherent model. In general, the pollen records so far available from monsoonal China, can only suggest that human disturbance becomes an increasingly important factor in vegetation during the last 2000yr. At most sites in monsoonal China, particularly in the north, the pollen evidence for human impacts on vegetation before 2 ka and how the anthropogenic signal can be distinguished from the climatic signal are still yet to be explored (Fig. 9; Liu and Qiu, 1994; Ren and Beug, 2002). More fossil Y. Zhao et al. / Earth-Science Reviews 97 (2009) 242–256 pollen records with reliable chronology at high sampling resolution are needed from monsoonal China to investigate the degree of human influence on natural vegetation. 5. Concluding remarks 1. Fossil pollen data show clear changes in natural vegetation during the Holocene in all vegetation regions that are affected by the summer monsoon, including tropical seasonal rain forest, subtropical forest, temperate deciduous forest, conifer–deciduous mixed forest, temperate steppe and highland meadow/steppe. Despite various vegetation sequences in different regions, our synthesis results show a generally humid climate during the early and middle Holocene and a drier climate during the late Holocene. 2. Effective moisture inferred from fossil pollen records in eastern monsoonal China correlates with the summer monsoon intensity pattern as indicated from other independent climate records. However, effective moisture inferred from fossil pollen records shows only a gradual increase at 11.5–9.5 ka, with ca. 1000-year delay after the precipitation maximum as indicated by monsoon intensity records from Dongge Cave and Sanbao Cave. Pollen-inferred moisture change shows a decreasing trend from 6 to 5 ka, with an abrupt shift at ca. 4.5 ka, probably due to the decrease in monsoon intensity. 3. Rate of change analysis of fossil pollen data shows that rapid vegetation change occurred at 11–10 ka, 6–5 ka and 2–1 ka, likely in response to shifts in summer monsoon strengths for the first two periods but more likely induced by human disturbance for the latest interval. 4. Human activities could be a very important factor affecting natural vegetation at large scale during the last 2 ka. More palynological and archaeological data will be needed to evaluate the relative contributions from climatic and anthropogenic factors to vegetation changes during the late Holocene. 5. High-resolution records with robust chronology for vegetation and climate reconstructions are still lacking, particularly in tropical monsoonal rain forest, temperate deciduous forest and conifer– deciduous mixed forest regions. Acknowledgements We thank the following individuals for providing the original pollen data: Shuyun Wang and Houyuan Lu (Huguangyan Maar Lake), Hongyan Liu (Haoluku), Ulrike Herzschuh (Zigetang Lake), Qinghai Xu and Jule Xiao (Daihai Lake), Ji Shen (Qinghai Lake), Wenying Jiang (Bayanchagan Lake), and Zhaodong Feng and Chengbang An (Dadiwan); and Yuting Hou and Xiaoli Guo for digitizing some pollen data. We thank two anonymous reviewers for their helpful comments and suggestions that improved the manuscript. This project was supported by the National Natural Science Foundation (NSFC grant no. 40771212), NSFC Innovation Team Project (no. 40721061) and MOE Program for New Century Excellent Talents in University. 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