Journal of Plant Ecology Volume 9, Number 1, Pages 51–60 February 2016 doi:10.1093/jpe/rtv029 Advance Access publication 18 March 2015 available online at www.jpe.oxfordjournals.org Growth–climate response and drought reconstruction from tree-ring of Mongolian pine in Hulunbuir, Northeast China Zhongjie Shi1,†, Lihong Xu2,†, Linshui Dong3, Jixi Gao4,*, Xiaohui Yang1, Shihai Lü5, Chaoyang Feng5, Jianxun Shang6, Aiyun Song3, Hao Guo1 and Xiao Zhang1 1 Institute of Desertification Studies, Chinese Academy of Forestry, 10 Huaishuju Road, Beijing 100091, China Institute of Forest Ecology, Environment and Protection, Chinese Academy of Forestry, 1 Dongxiaofu, Xiangshan Road, Beijing 100091, China 3 Key Laboratory of Eco-environmental Science for Yellow River Delta of Shandong Province, Binzhou University, 391 Huanghewu Road, Binzhou 256603, China 4 Nanjing Institute of Environmental Sciences, Ministry of Environmental Protection, 8 Jiangwangmiao Street, Nanjing 2100422, China 5 Chinese Research Academy of Environmental Sciences, 8 Dayangfang, Beiyuan Road, Beijing 100012, China 6 Songliao Water Resources Commission, Ministry of Water Resources, 4188 Jiefang Road, Changchun 130021, China *Correspondence address. Nanjing Institute of Environmental Sciences, Ministry of Environmental Protection, No. 8 Jiangwangmiao Street, Nanjing 210042, China. Tel: +86-25-85-28-72-90; Fax: +86-25-85-28-72-78; E-mail: [email protected] † These authors contributed equally to this work. 2 Abstract Aims Drought affected by atmosphere–ocean cycle is a dominant factor influencing tree radial growth of sandy Mongolian pine (Pinus sylvestris var. mongolica) and regional vegetation dynamics in Hulunbuir, China. However, historical droughts and its correlations with tree radial growth and atmosphere–ocean cycle in this area have been little tested. Methods We developed tree-ring chronologies of Mongolian pine from Hulunbuir, Inner Mongolia, China and analyzed the correlations between tree-ring width index, the normalized difference vegetation index and Palmer drought severity index (PDSI), then developed a linear model to reconstruct the drought variability from 1829 to 2009. Long-term trends and its linkages with atmosphere–ocean cycle were performed by the power spectral, wavelet and teleconnection analysis. Important Findings The local moisture variations affected largely the regional vegetation dynamics and tree-ring growth of Mongolia pine in the forest– grassland transition. Using tree-ring width chronology of Mongolian pine, the reconstruction explains 49.2% of PDSI variance during their common data period (1951–2005). The reconstruction gives a broad-scale regional representation of PDSI in the Hulunbuir area, with drought occurrences in the 1850s, 1900s, 1920s, mid-1930s and at the turn of the 21st century. Comparisons with other treering drought reconstructions and historical records reveal some common drought periods and drying trends in recent decades at the northern margin zones of the East Asian summer monsoon (EASM). The drying trends in these zones occurred earlier than weakening of the EASM. A REDFIT spectral analysis shows significant peaks at 7.2, 3.9, 2.7–2.8, 2.4 and 2.2 years with a 0.05 significance level, and 36.9, 18.1 and 5.0 years with 0.1 significance level. Wavelet analysis also shows similar cycles. Drought variations in the study area significantly correlated with sea surface temperatures in the western tropical Pacific Ocean and middle and northern Indian Ocean, and the Pacific Decadal Oscillation and North Atlantic Oscillation. This suggests a possible linkage with the El Niño-Southern Oscillation, the EASM and the Westerlies. Keywords: tree-rings, dendroclimatology, Mongolian pine, drought, PDSI, SSTs, East Asian summer monsoon (EASM), Hulunbuir, China Received: 22 May 2014, Revised: 9 February 2015, Accepted: 27 February 2015 © The Author 2015. Published by Oxford University Press on behalf of the Institute of Botany, Chinese Academy of Sciences and the Botanical Society of China. All rights reserved. For permissions, please email: [email protected] 52 INTRODUCTION The Hulunbuir area of northeast China is a transitional zone between semiarid and arid conditions, monsoon and non-monsoon climate, forest and steppe, and agricultural land and pastureland. Moisture in the area is affected by the Asian summer monsoon, and it is extremely sensitive and vulnerable to climatic and environmental changes (Fang et al. 2012). For example, a weakening of the Asian summer monsoon in northern China since the 1950s corresponds well with a drying tendency during the period (Jiang and Wang 2005; Xu et al. 2011). Severe drought events have significantly impacted local populations, particularly in arid and semiarid regions (Easter et al. 2000). For instance, one such event in the 1920s and 1930s in northern China caused widespread famine and tremendous loss of life (Liang et al. 2007). However, few reliable instrumental records extend back 100 years in China (Bradley 1999). The situation is even worse in northeast China, where weather records extend back only 60 years. The limited meteorological records from this area have hindered our understanding of the processes and mechanisms of past climate variability. Long-term records from tree-rings have been used to extend these limited meteorological records and help in detecting the dynamics of the Asian summer monsoon on longer timescales (Cook et al. 2010; D’Arrigo et al. 2000, 2001; Davi et al. 2009, 2010; Liu et al. 2009a; Mann et al. 2009; Pederson et al. 2001). Here, we use annual tree-ring width variation as a proxy for drought and for extending drought records in the Hulunbuir area. The Palmer drought severity index (PDSI) has been reconstructed for the Ortindag Sand Land (Liang et al. 2007), Guiqing Mountain (Fang et al. 2010), Kongtong Mountain (Fang et al. 2012), Qilian Mountains (Deng et al. 2013; Lu et al. 2013) and Guancen Mountain (Sun et al. 2012). The reconstructed drought variations have been used to explore the connection between drought and variability of the East Asian summer monsoon (EASM) on larger spatial and temporal scales (Liu et al. 2010). Recently, great progress has been made in dendroclimatological studies in the Inner Mongolia (Liang et al. 2008; Liu et al. 2007, 2010). In addition, annual precipitation and temperature have been reconstructed for the Hailar region (Bao et al. 2012; Liu et al. 2009b), Helan Mountain (Liu et al. 2005) and Xinglong Mountain (Liu et al. 2012). Nevertheless, drought reconstruction has seldom been done for the Hulunbuir area. The normalized difference vegetation index (NDVI) is an ideal indicator of vegetation activity such as forest and steppe, which has been used to study the photosynthesis, land surface evapotranspiration, exchange of CO2 and energy between the atmosphere and land surface (Chen et al. 2012). Recently, many studies had revealed the relationships between the tree-ring growth and NDVI from forest (Kaufmann et al. 2004; Leavitt et al. 2008) and grassland (He and Shao 2006; Liang et al. 2009) at different spatial and temporal scales. However, no attempt has been made to explore the relationship and mechanism between NDVI, tree-rings and drought in the transitional zone of forest and steppe. Journal of Plant Ecology The aims of this paper are (i) to test potential relationships between tree radial growth, NDVI and PDSI change, (ii) to construct regional drought variability based on the tree-ring record from Hulunbuir and (iii) to explore links between reconstructed drought records and the atmosphere–ocean cycle. MATERIALS AND METHODS Study area The study area is in Hulunbuir of eastern Inner Mongolia, an ecological transitional zone between steppe and forest, locating on the present margin of the EASM (Lee and Zhang 2011). The region has a typical temperate continental climate; it is hot and rainy in summer and cold and dry in winter, with annual average temperature of −2°C. Temperature in January averages −25.8°C and can reach a lowest value of −48.5°C, whereas in July, it averages 19.9°C and can reach a highest value of 36.7°C. Average annual precipitation is ~350 mm. This falls mainly from June through August, accounting for 60–70% of the annual amount. Annual evaporation is ~1210 mm, three times the rainfall. The main regional tree species is Mongolian pine (Pinus sylvestris var. mongolica). Tree-ring and NDVI data The tree-ring samples were collected from living trees in Xishan Forest Park (49°12ʹN, 119°42ʹE; 515–669 m above sea level) of Hulunbuir, during October 2009. Cores were taken from the trees by an increment borer at breast height, with two or three cores per tree. In total, 44 radii from 26 trees were used for drought reconstruction. Tree-ring samples were pre-treated according to the standard protocols of Stokes and Smiley (1968). After the cores were mounted, dried and sanded, we cross-dated (Fritts 1976) them using a skeleton diagram (Stokes and Smiley 1968) to ensure an accurate calendar year for each tree-ring. The width of each ring was measured with a LINTAB™ 5 ring analyzer (Frank Rinn, Germany) of accuracy 0.01 mm. Quality control of cross-dating was done using the COFECHA software program (Holmes 1983), and cores with any ambiguities (eight total) were eliminated. After cross-dating, individual tree-ring width measurements were detrended and standardized to the tree-ring width index, using the ARSTAN program (available at http:// www.ldeo.columbia.edu/tree-ring-laboratory; Holmes 1983). Undesirable growth trends related to tree age and stand dynamics, but unrelated to climatic variations, were removed from each series during the detrending process. To maximize the common signal at the lowest frequency possible and to avoid the impact of different growth rates at different ages, each tree-ring width sequence was standardized by fitting a negative exponential or straight line (Gonzalez-Elizondo et al. 2005; Liu et al. 2009b). All individual index sequences were combined into a single chronology by calculating the biweight robust mean, and standard (STD), residual (RES) Shi et al. | Tree-ring-based growth–climate response and drought reconstruction in Hulunbuir 53 and autoregressive (ARS) chronologies were determined. The STD chronology contains much lower frequency signals, and the RES chronology contains only common higher frequency signals. However, the ARS chronology includes both lower and higher common frequency signals of groups. The STD version of the chronology was used in further analysis, as it preserves much lower frequency signals (Cook and Kairiukstis 1990). The average percentage of absent rings in the samples was 0.02%. Subsample signal strength (SSS) was used to assess adequacy of replication in early years of the chronology (Wigley et al. 1984). To obtain the longest chronology and ensure reliability of reconstruction, the SSS of the first year of the chronology was restricted to at least 0.85. This threshold corresponds to a minimum sample depth of six trees beginning in 1829. NDVI, a normalized ratio of the near-infrared and red spectral reflections, was employed in this study as a proxy for vegetation growth and has been widely used in studies on vegetation growth at regional and global scales (Huber et al. 2011; Pettorelli et al. 2005; Piao et al. 2011). The data were derived from the NOAA/AVHRR Land data set (available at http://daac.gsfc. nasa.gov/) processed by the Global Inventory Monitoring and Modeling Studies group, with a spatial resolution of 8 × 8 km and had been collected at 15-day intervals (Tucker et al. 2005) from 1982 to 2006. These data were processed to account for orbital drift, to minimize cloud cover, to compensate for sensor degradation and sensor intercalibration differences and to consider the effects of stratospheric volcanic aerosols (Tucker et al. 2005). Using the 15-day NDVI data of each month, the monthly NDVI data were generated by the maximum value composite method and the annual and season NDVIs were generated separately by computing averages of the respective months. In this study, NDVI in the area (48–50°N, 118–120°E) were utilized. et al. 2007), we also investigated correlation between tree-ring width indices and regional PDSI data. PDSI is a meteorological drought metric based on a water balance model, in which temperature, precipitation and soil characteristics are considered (Dai et al. 2004; Palmer 1965). PDSI has been widely used in global. Positive and negative PDSI values correspond to wet and dry conditions, respectively. The PDSI herein was developed by Dai et al. (2004), and features a 2.5° × 2.5° gridding system. A monthly PDSI grid (48.75°N, 118.75°E) nearest our sites for 1951–2005 was used for the analysis. Statistical analysis Correlation analysis between the tree-ring width index and instrumental meteorological records was performed for the reliable time period of 1951–2005. And the correlations between NDVI and tree-ring width index/climatic variables were analyzed during 1982–2006. The PDSI reconstruction was conducted based on a split calibration-verification procedure designed to test model reliability (Fang et al. 2010). Statistics were used to evaluate model capability, including simple correlation coefficient (R), R-squared (R2), sign test (S), t-statistics, reduction of error (RE) and coefficient of efficiency (CE). Values of RE and CE > 0 indicate rigorous model skill (Cook et al. 1999). Teleconnections between regional droughts and remote oceans were analyzed by correlations between regional droughts and sea surface temperature (SST; Buckley et al. 2007; Li et al. 2008). We used the HadISST1 SST data set (Rayner et al. 2003), and spatial correlation was analyzed using the KNMI Climate Explorer (available at www.knmi. nl). Power spectral analysis was performed by PAleontological STatistics software (Hammer et al. 2001). Wavelet analysis was done using software available online (available at www. ion.researchsystems.com/IONScript/wavelet/; Torrence and Compo 1998). Climate data Precipitation and temperature data from nearby Hailar meteorological station (49°13ʹN, 119°45ʹE) were compared with the tree-ring width data for testing the climate–growth relationship. As some other studies in the moisture-sensitive area showed tree-ring width to be significantly correlated with PDSI in Northern China (Fang et al. 2009; Li et al. 2007; Liang RESULTS Chronology statistics Figure 1 shows the STD chronology and sample sizes during 1805–2009. Statistical characteristics of STD chronology are listed in Table 1. Of the 44 radii, 6 were >180 years old, Figure 1: STD ring-width index chronology and the number of samples for each year from Hailar. Thick line depicts 11-year moving average. 54 Journal of Plant Ecology Table 1: statistical characteristics of STD chronology Basic statistics STD First-order autocorrelation 0.41 Mean sensitivity 0.26 SD Signal-to-noise ratio EPS 0.28 35.81 0.97 Variance in first eigenvector 54.44% First year where SSS > 0.85 (number of trees) 1829 (6) while 21 were >150 years old. Mean tree-ring width was 0.99 ± 0.28 mm (mean ± standard deviation [SD]). Narrow rings mainly occurred in 1836, 1853, 1856, 1907, 1926, 1986, 1987, 2003, 2004, 2006, 2007 and 2008 (Fig. 1), which displayed an obvious growth decrease in the end of 20th and beginning of the 21st century. During the past 204 years, the occurrences of narrow rings were most frequent in the recent 30 years. More serious, the locally missing rings occurred in 1987. Mean sensitivity, a measure of relative difference in widths between adjacent rings, was 0.26. This indicates that tree growth was sensitive to changes of local environment. The first-order autocorrelation coefficient was 0.41, indicating that tree growth in a given year had an impact on growth the following year (Fritts 1976). The expressed population signal (EPS; Wigley et al. 1984), used to represent an acceptable level of chronology confidence, was 0.97, much higher than the EPS threshold 0.85. High variance in the first eigenvector (54.44%) and signal-to-noise ratio (35.81) indicates that the chronology contains considerable common signals and is thus suitable for climate change study. Tree-ring and NDVI series coherence Figure 2 showed the tree-ring width and NDVI series and their correlations during 1982–2006. The tree-ring width series corresponds well with the regional annual NDVI series (Fig. 2a). The tree-ring width series display high correlations with the regional NDVI series (Fig. 2b). The correlation coefficient reached up to highest value (r = 0.71, P < 0.01) in the monthly scale. The annual mean NDVI significantly correlated with tree-ring width (r = 0.43, P < 0.05) during 1982–2006. NDVI has significant correlations with the tree-ring widths during the growing season (May–September), especially during the period of May–July (r = 0.74, P < 0.01). The rapid growth of Mongolia pine occurred mainly in the period of May to September, especially during May to July (Zhao 1963). However, the accelerated growth for the steppe happened on the period of May to July. The vegetation on April was still in the dormant phase. A significant correlation was found between the tree-ring width and NDVI range of April to July (r = 0.58, P < 0.01). In addition, the spatial correlation between mean May–July NDVI and tree-ring widths also showed a higher correlation within the wide regions (see Supplementary Figure S1). The high correlations may be caused mainly by the similar phenological processes for the forest and steppe. These results also showed the coherence of regional vegetation dynamic from the satellite and tree-ring width from our observation. Tree-ring and NDVI response to climatic variables Correlation for the STD chronology with climatic variables from Hailar meteorological station (49°13ʹN, 119°45ʹE) and monthly PDSI (48.75°N, 118.75°E) is shown in Fig. 3. Treerings correlate negatively with mean temperature, and significant correlations at the 0.05 level were found in April and June–September of the current year (Fig. 3a). Tree-rings were significantly positively correlated with precipitation during May–August in the current year (P < 0.05), and the highest correlation coefficient was observed in July (Fig. 3a). Correlation with PDSI, a direct metric of moisture conditions (Dai et al. 2004), showed that tree radial growth was positively correlated with PDSI for all months between the previous October and the current September (Fig. 3b). This indicates that moisture availability during these seasons is the primary limiting factor to radial growth of Mongolia pine in the study area. The highest correlation of 0.70 is found in current May–July PDSI data. The study area is in a semiarid region, and annual precipitation cannot meet demands of tree evapotranspiration. Tree growth in this area is largely moisture-stressed. As a metric of soil moisture considering both precipitation and temperature, PDSI is a better indicator of moisture than simply precipitation. In addition, spatial correlation between the tree-ring index and the entire grid of the PDSI from Dai et al.(2004) showed that the tree-ring record reflects the average May–July PDSI over a sizable area around the sampling site (see Supplementary Figure S2). The climatic variables had an impact on the regional vegetation dynamics. NDVI in May was significantly and positively correlated with the precipitation and temperature. However, NDVI on June, July and August was found a significantly positive correlation with the precipitation on the previous month, respectively. In particular, the correlation analysis showed May–September NDVI was mainly influenced by the May–July precipitation. Compared with the above analysis, the higher correlations were displayed between NDVI and PDSI. NDVI in May, June and July was significantly correlated with their previous monthly PDSI, respectively (see Supplementary Table S1). The May–July NDVI was found a highest correlation with the concurrent PDSI (r = 0.74, P < 0.01). Above results displayed a similar climate response pattern for the regional NDVI and tree-ring growth of Mongolia pine, showing some common limiting factors for their growth. PDSI reconstruction and validation A simple linear regression model was used to reconstruct the PDSI from May to July of the current year: PDSI57 = 4.98*STD . –A ug –J ul . M ay . M ay . –A ug ep n. Ju l ua M ay –S ly A nn Ju ne Ju M ay Shi et al. | Tree-ring-based growth–climate response and drought reconstruction in Hulunbuir 55 Figure 2: tree-ring width and NDVI series (a), their correlations (b) from 1982 through 2006. Figure 3: correlation analyses of STD chronologies with monthly mean temperature and total monthly precipitation (a) and monthly PDSI (b). Dashed lines indicate 0.05 significance level. – 5.17, where PDSI57 is average May–July PDSI and STD represents the tree-ring STD chronology. The reconstructed PDSI agrees well with instrumental records (Fig. 4), explaining 49.2% of total variance during the calibration period 1951– 2005 (Table 2). To further evaluate statistical reliability of the reconstruction model, a split-period, calibration-verification procedure was used. As shown in Table 2, the calibration and verification results for the split sub-periods 1951–1978 and 1981–2005 generally show a good model fit. This is for the two most rigorous tests of model validation, RE and CE. These tests demonstrate the validity of the regression model. Drought variability The May–July PDSI, reconstructed from 1829 to 2009 for the Hulunbuir area, has mean value −0.25 and SD = ±1.42. This accords with the normal moisture level (PDSI = 0.0 ± 0.5) in the central and western Great Plains of the USA (Palmer 1965). In this study, the value of −0.25 was considered as the normal moisture condition, and years with values greater than twice the SD were defined as extreme dry or wet years. During the 181 years of reconstructed PDSI (1829–2009), extremely dry years were 1853, 1856, 1987 and 2003, and extremely wet years were 1934 and 1984. There is greater frequency of severe dry years during recent decades, especially after ~1986. We defined years with PDSI >1 SD above the mean as wet years, and those with PDSI <1 SD below the mean as dry years. The full PDSI reconstruction displays strong interannual variation across the entire period (Fig. 5). There were 28 dry years and 32 wet years in the reconstructed series, accounting for 15.47% and 17.68% of the total, respectively. Dry periods of 2 years or more were found during 1905–09, 1926–28, 1950–51, 1986–87, 2001–03 and 2006–08. Wet periods exceeding 2 years were during 1846– 48, 1868–69, 1881–83, 1932–34, 1939–41, 1953–56, 1983–84 and 1989–91. The PDSI reconstruction from the Monsoon Asian Drought Atlas (Cook et al. 2010) shows trends similar to our reconstruction (see Supplementary Figure S4). Droughts are shown in the 1850s, 1900s, 1920s, mid-1930s and turn of the 21st century. On the decadal timescale, the reconstructed PDSI series also reveals drought variability. An 11-year moving average of reconstructed PDSI (Fig. 5) shows distinct alternations of dry and wet intervals. There were six dry intervals with PDSI below the mean: 1837–39, 1851–62, 1889–1912, 1920–32, 1963–74 and 1997–2009. There were six wet intervals with PDSI above the mean: 1829–36, 1840–50, 1863–88, 1913– 19, 1933–62 and 1975–96. The longest wet period lasted for 30 years, from 1933 to 1962, and the longest dry period was 24 years, from 1889 to 1912. Figure 5 also demonstrates that after the 1940s, the reconstructed PDSI shows a longterm descending trend. This trend is more significant after the 1980s. Significant (95% confidence level) spectral peaks were revealed at 7.2, 3.9, 2.7–2.8, 2.4 and 2.2 years (Fig. 6) by the REDFIT spectral analysis (Hammer et al. 2001). We also found periods of 36.9, 18.1 and 5.0 years at a 90% confidence level. 56 Journal of Plant Ecology Wavelet analysis showed similar cycles to those found in our reconstruction by power spectral analysis (Fig. 7). To some extent, the power resided within the typical El Niño-Southern Oscillation (ENSO) variability bands during the entire reconstruction. The greatest power at decadal timescale, expressed in the range from 30 to 40 years, was from the 1950s to the turn of the 21st century. Power in the 10–20 year range was found from the 1840s–60s and 1890s–1910s. Power within the 8–12 and 4–8 year ranges was in the 1920s–40s and 1980s–90s (Fig. 7). DISCUSSION Drought–growth relationships Previous studies in arid and semiarid areas of North China have shown that tree growth is primarily determined by effective moisture (Fang et al. 2009, 2010; Li et al. 2007; Liang et al. 2007). The moisture-sensitive growth pattern is apparent from negative correlations with temperature and positive correlations with precipitation. Temperature affects tree growth via soil moisture amount, and this is supported by the negative correlations. The reconstruction agrees with this pattern. But compared to precipitation, temperature is more significant in causing drought stress as demonstrated by higher coefficients, which is also evident at Guiqing Mountain (Fang et al. 2010), Kongtong Mountain (Fang et al. 2012), Guancen Mountain (Sun et al. 2012), and Ortindag Sand Land (Liang et al. 2007). During the growing season, higher temperature can accelerate tree transpiration and respiration, simultaneously decrease carbohydrate accumulation in stems. Annual effective evapotranspiration is about five to seven times greater than total precipitation (Bao et al. 2012), which means that high temperature would exacerbate the large soil moisture deficits. Temperatureinduced drought may be common in extremely dry regions or locations with sandy soil, where potential evapotranspiration is considerably higher than actual evapotranspiration. For such areas, drought-growth correlations are expected to be high, which may account for the high PDSI–growth correlations in the Hulunbuir area. These findings are similar to results of studies of the Qinghai-Tibet Plateau (Gou et al. 2008; Qin et al. 2008; Shi et al. 2010), north-central Shaanxi Province (Cai et al. 2008), Lüliang Mountains (Cai et al. 2010), and Funiu Mountains (Tian et al. 2009). The occurrence of missing or narrow rings indicates that drought stress sometimes controls its survival limit (Liang et al. 2014). The dry and warm year appear to cause the lower tree-ring growth, even missing rings. In the past 30 years, 7 years with the narrower rings were consistent with the droughts in the study area. In particular, the drought in 1987 was the most serious in the past 181 years, the locally missing rings occurred. Similar to the relationship between tree-ring width and drought, the lower NDVI values also coincide with the drought events. The high correlations were observed among the ring width, NDVI and PDSI, particularly in the growing season, which appears to be a common feature of the response of the tree growth, regional vegetation on the climate for the grassland and forest in the semiarid and arid regions of northern China. The significant linkage between NDVI and ring growth is a result of their common phenological processes. It is more important that there is a common and similar influence of seasonal moisture on the growth of different vegetation types in the study area. The PDSI data here faithfully represent local drought conditions, and strongly correlated with tree growth and regional vegetation dynamics. The most significant moisture limitation is during May and June, which have relatively high temperature and light precipitation (Fig. 3b) and indicate a greater soil moisture limitation. Drought variability Figure 4: comparison of observed and reconstructed May–July PDSI from 1951 to 2005. The 1920s–30s drought was reported in other studies of arid and semiarid regions in North China (Fang et al. 2009; Liang Table 2: statistics of calibration and verification tests of the PDSI57 reconstruction Calibration Period Verification r r2 S T Period r RE CE S T 1981–2005 0.73 0.54 19/6* 4.25 1951–1980 0.68 0.65 0.40 26/4** 4.61 1951–1978 0.75 0.56 26/2** 4.38 1979–2005 0.69 0.29 0.25 20/7* 4.27 1951–2005 0.70 0.49 45/10** 6.09 — — — — — — *P < 0.05, **P < 0.01. Shi et al. | Tree-ring-based growth–climate response and drought reconstruction in Hulunbuir 57 Figure 5: reconstructed PDSI and 11-year smoothed average reconstruction. Figure 6: spectral density of reconstructed PDSI during 1829–2009 in Hulunbuir area. Spectral analysis is based on REDFIT method. et al. 2006; Liu et al. 2005, 2009b, 2010; Yang et al. 2011; Zhang et al. 2011). This drought was widespread and varied in intensity and length across this region, leading to tremendous loss of life and severe damage to the economy. The dry epoch after 1986, especially after 1995, was the most severe over the past 181 years in the Hulunbuir drought reconstruction. This is consistent with an observed drying trend since the 1980s (Deng et al. 2013; Fang et al. 2009, 2010, 2012; Li et al. 2007; Shi et al. 2007), which was also recorded at Xinglong Mountain (Fang et al. 2009; Liu et al. 2013), Guiqing Mountain (Fang et al. 2010), Kongtong Mountain (Fang et al. 2012) and Qilian Mountain (Deng et al. 2013; Yang et al. 2011). This drought has increased the risk of water resource and ecosystem vulnerability, especially in our study area. The droughts occurred in 1987 and 2003 were the most severe events during 1829–2009, and were centered on an area similar to that for which our tree-rings are representative (see Supplementary Figure S3), strongly corresponding to warm phases of ENSO. In addition, the wet intervals during 1863–88 and 1933–62 also correspond to the longest wet intervals from a tree-ring-based PDSI reconstruction in the Ortindag Sand Land (Liang et al. 2007). Comparison with other studies A severe drought in North China from 1904 to 1909 was captured by the Hulunbuir PDSI reconstruction. According to historical records (Ding 2008), the drought began in western Heilongjiang Province (the Hulunbuir area of today) in 1904. Figure 7: wavelet plots for reconstructed May–July PDSI. Areas surrounded by thick black contours (5% significant level, using rednoise [ARS lag 1] background spectrum) are regions of significant power at corresponding timescales. The drought was continuous thereafter, and there was no rainfall from spring to autumn of 1909. Only 30–40% of crops compared with a normal harvest year were harvested during this period because of severe summer drought. This illustrates the reliability of the sequence reconstructed here. In the mid-late 1920s, an extreme drought disaster caused serious social and economic problems in northern China. In 1926, April–July precipitation was 50% lower than the average in Hailar. The drought recurred in summer 1927, and its continuation prevented crop harvesting in 1928 (Ding 2008; Shen 2008). Moreover, in 1935, most areas around Hulunbuir were subjected to continuous drought in spring and summer; this drought was especially severe in Hailar (Shen 2008). Such continuous drought agrees with results of tree-ring temperature reconstruction in many regions of northern China (Cai et al. 2008; Liang et al. 2007). According to historical records, the continuous severe drought between 1876 and 1878 represents one of the greatest natural hazards during the Qing Dynasty in middle and northern China. In the latter region, this event affected 1.6–2 billion people (~50% of the total Chinese population) and led to 10 million fatalities (Li et al. 1994; Shen 2008). However, this extreme drought was not evident in the Hulunbuir reconstruction, which shows a normal state during this period. 58 We also compared the reconstructed drought variation with seven other tree-ring-based PDSI reconstructions near our sampling sites (see Supplementary Figure S4; Cook et al. 2010; Deng et al. 2013; Fang et al. 2010, 2012; Liang et al. 2007). Our reconstruction shows drought intervals similar to the other seven drought reconstructions (see Supplementary Figure S4). Over the past 181 years, there were common climate trends among the various series, including the wet period of the 1860s–70s (see Supplementary Figure S4, period I), drought periods of the 1920s–30s (see Supplementary Figure S4, period II) and the turn of the 20th century (except at Guiqing Mountain; see Supplementary Figure S4, period III). These reconstructions also show a slow drying trend in recent decades (see Supplementary Figure S4 a~h). By comparing these trends at different sites, we found that the drying trend started at the end of the 1930s at Hulunbuir (Cook et al., 2010; this study) and Helan Mountain (Fang et al. 2010), and at Ortindag Sand Land (Liang et al. 2007) and the eastern Qilian Mountains (Deng et al. 2013) since the 1950s. This trend began in the 1970s at Kongtong Mountain (Fang et al. 2012) and Xinglong Mountain (Fang et al. 2010), and in the 1980s at Guiqing Mountain (Fang et al. 2010; see Supplementary Figure S4). The eight studies (including this study) were done in the transitional zones of arid and semiarid climate, and at the margin of the EASM. Despite some differences, this comparison demonstrates a spatial and temporal link to PDSI in these environmentally sensitive EASM margin zones. In addition, the common trends within the eight PDSI time series reveal a drying trend during the second half of the 20th century in these regions, and this trend is consistent with weakening of the EASM (Jiang and Wang 2005; Liu et al. 2003). Another study has shown that EASM has been weakening since the 1950s (Jiang and Wang 2005; Xu et al. 2011), and sites within marginal monsoon zones experienced this weakening earlier than in monsoon zones. In addition, the weakening in the northern marginal monsoon zones occurred earlier than the south marginal zones. Linkages with remote oceans The 2–4-year peak cycle found in some moisture-related reconstructions by tree-ring data in North China (Deng et al. 2013; Fang et al. 2009, 2010, 2012; Liu et al. 2009b) falls within the range of ENSO variability (Allan and Parker 1996), suggesting a link between droughts and ENSO. And the link was supported by the negative correlation between the PDSI and western tropical Pacific SSTs (see Supplementary Figure S5), i.e. the higher SSTs induced the occurrence of drought in Hulunbuir area. Spatial correlation between PDSI and SST within the instrumental period (1951–2005) also displayed patterns similar to those of Guiqing Mountain (Fang et al. 2010), the Qilian Mountains (Deng et al. 2013) and northeastern Tibetan Plateau (Li et al. 2008). Some studies in northeast China have shown a climate response of EASM to moisture advection from the western tropical Pacific (Huang et al. 2007; Journal of Plant Ecology Sun et al. 2007). This suggests a teleconnection between local droughts and the EASM. Drought variability was greatly affected by the Pacific Decadal Oscillation (PDO). On an annual scale, our reconstructed PDSI correlates with May–July PDO for 1900–2009 (r = 0.122, P < 0.2). On a decadal scale, this PDSI also significantly correlated with May–July PDO (r = 0.257, P < 0.01). Prevailing cool PDO regimes from 1920 to 1932 and 1963 to 1974, and warm PDO regimes from 1933 to 1962 and 1975 to the mid-1990s (Mantua and Hare 2002), correspond with most of our dry and wet periods. Another study showed that the Hulunbuir area is within the zone of the Westerlies (Lee and Zhang 2011). There is significant correlation (r = 0.189, P < 0.05) between our reconstructed PDSI and the North Atlantic Oscillation (NAO), which is an indicator of the strength of these Westerlies. This indicates their possible influence, simultaneous with the period of EASM decline. This influence was also found in the Qilian Mountains within the EASM margin zone (Deng et al. 2013; Zhang et al. 2009) and Northwest China (Lee and Zhang 2011). CONCLUSIONS A tree-ring chronology from Mongolia pine spanning 1829– 2009 was developed for the Hulunbuir area of Northeast China. The PDSI was reconstructed using a model that explained 49.2% of variance. Spatial correlation analyses highlighted the broad-scale regional representativeness of our reconstruction. The moisture-sensitive growth pattern showed negative correlations with temperature and positive correlations with precipitation in the arid or semiarid region. The local moisture variations affected largely the regional vegetation dynamics and tree-ring growth of Mongolia pine in the forest–grassland transition. The strong PDSI–growth relationship demonstrates a capability for modeling local drought conditions. Drought reconstruction showed strong interannual and decadal variability, and drought events were revealed in the 1850s, 1900s, 1920s, mid-1930s and at the turn of the 21st century. Comparisons with other tree-ring drought reconstructions and certain historical records, they disclose jointly some common drought periods and drying trends in northern margin zones of the EASM. The latter drying trends preceded EASM weakening. The REDFIT spectral analysis showed significant peaks at 7.2, 3.9, 2.7–2.8, 2.4 and 2.2 years with 95% confidence level, and 36.9, 18.1 and 5.0 years with 90% confidence level. The wavelet analysis showed cycles similar to those in the reconstruction by power spectral analysis. Further, drought variations in our reconstruction significantly correlated with SSTs in the western tropical Pacific, mid and northern Indian Ocean, the PDO and NAO. These suggest links to ENSO, EASM and the Westerlies. SUPPLEMENTARY DATA Supplementary material is available at Journal of Plant Ecology online. Shi et al. | Tree-ring-based growth–climate response and drought reconstruction in Hulunbuir 59 FUNDING ‘948’ Project of State Forestry Administration China (20154-27), International S&T Cooperation Program of China (2015DFR31130), National Natural Science Foundation of China (41271033, 41471029 and 41371500) and The Lecture and Study Program for Outstanding Scholars from Home and Abroad (CAFYBB2011007). Davi N, Jacoby G, Fang K, et al. (2010) Reconstructing drought variability for Mongolia based on a large‐scale tree ring network: 1520–1993. J Geophys Res 115:D22103. Davi NK, Jacoby GC, D’Arrigo RD, et al. (2009) A tree ring-based drought index reconstruction for far-western Mongolia 1565– 2004. Int J Climatol 29:1508–14. Deng Y, Gou XH, Gao LL, et al. (2013) Aridity changes in the eastern Qilian Mountains since AD 1856 reconstructed from tree-rings. Quatern Int 283:78–84. 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