Tree-ring growth of Gmelin larch under contrasting local conditions in the north of Central Siberia

While the forest-tundra zone in Siberia, Russia has been dendroclimatologically well-studied in recent decades, much less emphasis has been given to a wide belt of northern taiga larch forests located to the south. In this study, climate and local site conditions are explored to trace their influence on radial growth of Gmelin larch (Larix gmelinii (Rupr.) Rupr.) trees developed on permafrost soils in the northern taiga. Three dendrochronological sites characterized by great differences in thermo-hydrological regime of soils were established along a short (ca. 100 m long) transect: on a river bank (RB), at riparian zone of a stream (RZ) and on a terrace (TER). Comparative analysis of the rate and year-to-year dynamics of tree radial growth among sites revealed considerable difference in both raw and standardized tree-ring width (TRW) chronologies obtained for the RZ site, characterized by shallow soil active layer depth and saturated soils. Results of dendroclimatic analysis indicated that tree-ring growth at all the sites is mostly defined by climatic conditions of a previous year and precipitation has stronger effect on TRW chronologies in comparison to the air temperatures. Remarkably, a great difference in the climatic response of TRW chronologies has been obtained for trees growing within a very short distance from each other. The positive relation of tree-ring growth with precipitation, and negative to temperature was observed in the dry site RB. In contrary, precipitation negatively and temperature positively influenced tree radial growth of larch at the water saturated RZ. Thus, a complicate response of northern Siberian larch forest productivity to the possible climate changes is expected due to great mosaic of site conditions and variability of environmental factors controlling tree-ring growth at different sites. Our study demonstrates the new possibilities for the future dendroclimatic research in the region, as various climatic parameters can be reconstructed from tree-ring chronologies obtained for different sites.     

Responses to climate change in radial growth of Picea schrenkiana along elevations of the eastern Tianshan Mountains,northwest China

Tree growth is largely driven by climate conditions in arid and alpine areas. A strong change in climate from warm-dry to warm-wet has already been observed in northwest China. However, little is known about the impacts of regional climate variability on the radial growth of trees along elevations of the eastern Tianshan Mountains. Consequently, we developed three tree-ring width chronologies of Schrenk spruce (Picea schrenkiana Fisch. et Mey.) ranging in elevation from 2159 to 2552 m above sea level (a.s.l.), which play an important role in the forestry ecosystem, agriculture, and local economy of Central Asia. In our study, the correlation analyses of growth-drought using the monthly standardized precipitation-evapotranspiration index (SPEI) at different temporal scales demonstrated that drought in growing season was the main factor limiting tree growth, regardless of elevation. The relationships between radial growth of Schrenk spruce and main climate factors were relatively stable by moving correlation function, and the trend of STD chronologies and basal area increment (BAI) also showed a synchronous decline across the three elevations in recent decades. And meanwhile, slight differences in responses to climate change in radial growth along elevations were examined. The drought stress increased as elevations decreased. Radial growth at the higher elevation depended on moisture availability due to high temperature, as indicated by the significant negative correlation with mean temperature in the late growing season of the previous year (August-September, p < 0.001). However, radial growth at the lower elevation were restricted by drought stress due to less precipitation and higher temperatures, as demonstrated by the significant negative correlation with mean temperature but positive with total precipitation in the early growing season of the current year (April-May, p < 0.05). In addition, the decline of radial growth (BAI) at the higher elevation (3.710 cm² yr⁻¹/decade, p < 0.001) was faster than that of the middle elevation (2.344 cm² yr⁻¹/decade, p < 0.001) and the lower elevation (3.005 cm² yr⁻¹/decade, p < 0.001) since 2000, indicating that the trees at higher elevation of a relatively humid environment were more susceptible to the effects of climate change due to their poor adaptability to water deficit. Therefore, the forest ecosystems would be suppressed as a result of increasing drought stress in the future, especially in the high-elevation forests of arid and semi-arid areas.     

Evaluating the temperature sensitivity of radial growth patterns from whitebark pine in the western Canadian Cordillera

We present a network of thirteen annual ring-width chronologies from high elevation whitebark pine (Pinus albicaulis Engelm.) sites in the western Canadian Cordillera in order to assess the dendroclimatic potential of this long-lived tree species. The temperature signal within the chronologies is complex and strongly influenced by diverging trends in the summer temperature and ring-width records from across the region. A first differences transformation of the tree-ring and temperature records illustrates a loss of frequency coherence in growth response to summer temperatures following reduced radial growth in the 1950s. Prior to reduced growth, we note a positive association with summer temperatures for both first differenced (r_d = 0.60) and traditional (r = 0.50) records. Following reduced growth, the association at first differences is maintained (r_d = 0.49) whereas there is a change in the lower frequency component of tree growth response to summer temperatures (r = ?0.34). We suggest the cause of this reduced temperature sensitivity is related to the interaction between diurnal temperature and cloud cover patterns, the hydrological regime of snowpack, and site conditions which have been amenable to the initiation of moisture stress during the latter half of the 20th century. Reduced radial growth is coincident with the arrival of white pine blister rust (Cronatium ribicola J.C. Fisch. ex Raben) into the study region which suggests this infestation may be related to the observed reduction in radial growth. Whitebark pine has considerable potential for the field of dendroclimatology. Unfortunately, the decline of the species due to the combined effects of climate change, white pine blister rust, mountain pine beetle (Dendroctonus ponderosae Hopk.), and forest fire exclusion practices indicate this potential may remain unfulfilled.     

Age-dependent tree-ring growth responses of Schrenk spruce (Picea schrenkiana) to climate—A case study in the Tianshan Mountain,China

For both its climatic and ecological importance, Schrenk spruce (Picea schrenkiana) is a crucial tree species living at mid-altitude on the western area of the Tianshan Mountains. It plays a key role on understanding climatic change in the Tianshan Mountains in the past 500 years. However, whether the relationship between tree growth and limiting climate factors is stable over time is still not well-known. In this study, standard and residual chronologies of four 100-year age classes (AC1 < 110a, 110a < AC2 < 210a, 210a < AC3 < 310a and AC4 > 310a) were established for detecting divergence in climate–growth relationships as well as comparing low-frequency and high-frequency variations. The results show that climate can account for a high amount of variance in tree-ring width and higher climate sensitivity was detected in younger trees. Younger trees (<210a) exhibit significantly negative growth responses to mean monthly air temperature of previous June and positive relationship with total monthly precipitation of current April and May, while mean monthly air temperature of current March may inhibit growth of older trees (>210a). Tree-ring chronology statistics and response function reveal that the age-growth patterns are non-monotonic. Our results together with previous studies demonstrate that the age effects on tree-ring growth–climate response is attributed to a combination of genetic characteristics and site microclimate, which suggests that it is necessary to consider both age-dependent and species-specific climate responses when using tree-ring measurements as a proxy for valid climate reconstructions.     

A 1700-year Athrotaxis selaginoides tree-ring width chronology from southeastern Australia

Few Southern Hemisphere tree-ring chronologies exceed 1000 years in length. We present a ca. 1700 years of indexed values for the long-lived conifer Athrotaxis selaginoides at Cradle Mt in southeastern Australia and compare it with the only other published millennial-plus length tree-ring chronology for Australia: the nearby Mt Read Lagarostrobos franklinii. We use simple correlation function and pointer year analyses to compare the climate responses of the two species (temperature, precipitation and growing degree days). Both chronologies show accelerated growth at their modern ends, but this growth acceleration is not synchronous, beginning approximately a quarter of a century earlier at the Cradle Mt site. This discrepancy may highlight the relevance of chronology composition and/or physiological differences in the species. Although the seasonality of the climatic responses of the two species is similar, that of A. selaginoides is generally weaker than that of L. franklinii. Somewhat paradoxically, the only pointer years in common between the chronologies are 1898 and 1908 CE. The periods from 600 to 900 CE and ∼1200–1450 CE are conspicuous for their absence of positive pointer years while no negative pointer years occur for either site from ∼1200–1350 CE. It is possible that differing patterns of pointer years can be partially explained by a peak in establishment from ∼1150–1850 CE at the Mt Read L. franklinii site compared to continuous establishment at Cradle Mt. Although statistically significant and time-stable climate responses for the A. selaginoides chronology are too weak to base a single-chronology climate reconstruction on, the long chronology will likely make an important contribution to future multi-proxy temperature reconstructions for southeastern Australia.     

The temperature sensitivity along elevational gradients is more stable in maximum latewood density than tree-ring width

Tree ring-based temperature reconstructions are preferably derived from maximum latewood density (MXD) compared to tree-ring width (TRW). Although temperature signals in MXD are less dependent on site ecology, systematic analyses of the effects of elevation and slope aspect on ring formation are still lacking. Here, we assess the climate sensitivity of MXD and TRW chronologies from six larch (Larix decidua Mill.) sites across the Simplon valley in the southwestern Swiss Alps, representing elevations from 1400 to 2150 m asl on both north- and south-facing slopes. We find decreasing temperature signals with decreasing elevation in MXD and TRW, though correlation coefficients are generally higher for MXD and on the warmer and dryer south exposed slopes. While the greatest temperature signals are found for MJJA at highest elevations with r = 0.71 for MXD and r = 0.57 for TRW (both p < 0.05 and for the 1928–2009 common period), MXD still correlates significantly positive at the lowest elevation site that is ~750 m below the treeline. Our findings indicate the suitability of MXD over TRW for temperature reconstructions when using historical wood sources of unknown origin.     

Process-based modeling of tree-ring formation and its relationships with climate on the Tibetan Plateau

Response of climate warming on tree-ring formation has attracted much attention during recent years. However, most studies are based on statistical analysis, lacking understanding of tree-physiological processes, especially in the mountainous regions of the Tibetan Plateau (TP). Herein, we firstly use an updated new version of the tree-ring process-based Vaganov-Shashkin model (VS-oscilloscope) to simulate tree-ring formation and its relationships with climate factors during the past six decades. Our analyses covered 341 sampled trees growing within elevations ranging from 2750 to 4575 m a.s.l. at five sampling sites across the TP. Simulated tree-ring width series are significantly (p < 0.01) correlated with actual tree-ring width chronologies during their common interval periods. Starting dates of tree-ring formation are determined by temperature at all five sampling sites. After the initiation of tree stem cambial activity, soil moisture content has a significant effect on tree radial growth. Ending dates of cambial activity are driven by temperature over the whole study region. Simulated results indicate differences between wide and narrow tree-rings are mostly induced by soil moisture content, especially during the first half of the growing season, when effects from temperature variations are minor. Interestingly, we detected significantly (p < 0.001) increased relative growth rates due to higher soil moisture content after the year 1985 at the five sampling sites. However, the variability of mean relative growth rates due to temperature is negligible before and after that. Based on the successful application of VS-oscilloscope modeling on the high-elevation tree stands on the TP, our study provides a new perspective on tree radial growth process and their varying relationships to climate factors during the past six decades.     

High-elevation inter-site differences in Mount Smolikas tree-ring width data

We present the longest high-elevation tree-ring width dataset in the Mediterranean reaching back to the 6th century CE. The network includes 101 living and 92 relict Pinus heldreichii Christ trees from four differently exposed sites in the 2100–2200 m a.s.l. elevation range of Mt. Smolikas in the Pindus Mountains in Greece. Though the sites were all sampled within a distance of <1 km, inter-site correlations are surprisingly low (r_1550–2014 = 0.65–0.87), indicating site exposure might affect tree-ring formation. We here explore the consequence of exposure differences on the climate signals in an eastern Mediterranean treeline ecotone. Temporally stable growth/climate relationships reveal similar seasonal patterns among the four sites, but differences in signal strength. P. heldreichii growth at Mt. Smolikas is significantly controlled by temperature in April (r_1951–2014 = 0.33–0.50) and precipitation in June-July (r_1951–2014 = 0.23–0.42), which emphasizes the overall importance of an early growth onset and subsequent moisture conditions. The association between stem growth and April climate is strongest in the South-facing stand, supporting the significance of higher insolation rates at this thermally privileged site. Strongest summer precipitation signals are found in the NE-facing stand, where trees seem to benefit least from an early growth onset and where reduced meltwater supply may enhance the dependency on early summer precipitation. The significance of spring temperature on tree growth in all four sites constrains the emergence of a distinct summer precipitation signal in the Mt. Smolikas high elevation ecotone. Exploration of the site-specific influences on a new millennium-long tree-ring width dataset is an important step towards an improved understanding of long-term climate variability in the Eastern Mediterranean. Site-related differences in climate sensitivity in the high-elevation tree-ring network at Mt. Smolikas indicate that both temperature and precipitation during different seasons could potentially be reconstructed if distinct site exposures (S versus NE) are considered.     

Elevation-specific tree-ring chronologies of Norway spruce and Silver fir in Southern Germany

Tree-ring (TR) chronologies are important instruments for the dating and provenance analyses of historical wood, as well as for climate reconstructions. However, radial growth patterns differ between tree species and growing environments. Therefore chronologies are more or less specific for a certain tree species, region and elevation. Chronologies that are restricted to more confined regions could extend the possibilities for dating, dendroprovenancing and regional climate reconstructions.In Southern Germany, the transport of wood by raft – for the supply of towns and cities with timber – has been documented since early mediaeval times. Consequently, not only local timber from the lowlands was used for construction purposes, but also alpine wood originating from sites up to the timberline. Since pronounced altitudinal gradients cause distinct climate differences, elevation-specific chronologies have the potential to improve dating precision in this region.In this contribution, a model is presented and applied in order to separate elevation-specific provenances of Norway spruce (Picea abies L. Karst) and Silver fir (Abies alba Mill.) in Southern Germany. The model is derived from more than 2100 living TR series originating from sites between 200 and 1710 m above see level. Absolute (mean, maximum) ring-width values and the variation of ring-widths (mean sensitivity) show distinct correlations with their altitudinal provenance, reflecting improvements in growth conditions as well as the increase of their yearly variation from high over intermediate to low elevation sites. Mean ring widths and mean sensitivity values were used as independent parameters in an exponential regression model which exhibits a coefficient of determination (r²) of 77% for spruce and 74% for fir. The prediction accuracy of the elevation amounts ±300 m for spruce and ±200 m for fir within the 95% confidence interval.The model was used to estimate the elevation origin of around 5000 historical spruce and 800 historical fir series from buildings located in Southern Germany. The historical TR series covering the AD 990–1800 period were allocated by the model to elevations between 120 and 2090 m. In a second step the individual TR series were combined to elevation-specific chronologies representing low, intermediate, and high altitudinal belts. The chronologies show distinct differences among the altitudinal belts in terms of signatures and pointer years, especially for spruce. Elevation-specific chronologies are assumed to amplify the dating possibilities of ancient timber, to provide valuable evidences for the origin of historical wood, and to offer more specific proxy data for regional climate reconstructions.     

Growth response of Sabina tibetica to climate factors along an elevation gradient in south Tibet

We examine the climate significance in tree-ring chronologies retrieved from Sabina tibetica Kom. (Tibetan juniper) at two sites ranging in elevation from 4124 to 4693 m above sea level (a.s.l.) in the Namling region, south Tibet. The study region is under the control of semi-arid plateau temperate climate. The samples were grouped into high- and low-elevation classes and standard ring-width chronologies for both classes were developed. Statistical analysis revealed a decreasing growth rate yet increasing chronology reliability with increasing elevation. Overall, correlation analyses showed that radial growth in S. tibetica at the study sites was controlled by similar climatic factors, regardless of elevation; these factors comprised early winter (November) and early summer (May–June) temperatures as well as annual precipitation (July–June). Slight differences in the correlation between tree growth along the elevation gradient and climate variables were examined. The correlations with early winter temperature varied from significantly positive at the low-elevation site to weakly positive at the high-elevation site, whereas the correlations between radial growth and early summer temperature increased from weakly negative at the low-elevation sites to strongly negative at the high-elevation sites. The abundant precipitation through the year may have masked variations in tree growth on different elevation aspects. Our results will aid future dendroclimatological studies of Namling tree rings in south Tibet and demonstrate the potential of S. tibetica Kom. for improving our understanding of environmental impacts on tree growth.     

Evidence for temperature limitation of nitrogen mineralisation in the Drakensberg Alpine Centre

We made use of pot experiments and soil mineralisation assays to test the effect of temperature on the soil nitrogen (N) economy of the Drakensberg Alpine Centre (‘mountain site’). The approach was enhanced by the inclusion of a contrasting warm, subtropical environment on the east coast of southern Africa (‘coast site’) which presented an opportunity to test plant growth in mountain soil outside of the mountain site's natural climatic envelope. This study was further augmented by two greenhouse experiments that helped isolate the factors responsible for the growth responses in the experiments above. Plant morphology, plant nutrients and soil nutrients were used as the basis for comparing treatment effects. The primary pot experiment showed that plant growth was uniform in the mountain site regardless of whether the test species was grown in intrinsically N-rich mountain soil or intrinsically N-poor coast soil. However, we noted significant growth differences at the coast site using the aforementioned soil nutrient regimes. In terms of the soil mineralisation assay, coast soil, derived from intrinsically N-poor sandstone, predictably mineralised little soil inorganic N at the mean spring temperature of 19 °C. However against expectations, the intrinsically N-rich mountain soil mineralised < 1% of its total soil N budget into inorganic N at 12 °C, most probably because the microbes responsible for the conversion of organic soil N to inorganic soil N were severely inhibited at this mean spring temperature. However, the potential to mineralise far more N in mountain soil was apparent when using an elevated experimental temperature of 30 °C, with 369% more soil N being available under the latter regime. Our results suggest that the cooler temperatures associated with high elevations in the mountain site constrain the activity of soil microbes in mountain soil, resulting in a functionally N-poor soil economy particularly deficient in inorganic N. This also explains the similar growth responses regardless of the soil being intrinsically N-rich or N-poor. We speculate whether or not more soil inorganic N may become available under a regime of warming due to accelerated N mineralisation, to the detriment of plant taxa adapted to low soil N availability.     

Climate-growth relationships for Rocky Mountain juniper (Juniperus scopulorum Sarg.) on the volcanic badlands of western New Mexico,USA

We sampled Rocky Mountain junipers (RMJ) to produce a multi-century tree-ring chronology from a relict lava flow, the Paxton Springs Malpais (PAX), in the Zuni Mountains of western New Mexico. Our objective was to assess crossdating potential for RMJ growing on the volcanic badlands of the region, investigate potential relationships between climate and RMJ growth, and investigate temporal variability in relationships identified between climate and RMJ growing at our site. We hypothesized that, similar to other drought stressed-conifers growing on the lava flows, RMJ responds to climate factors that influence and indicate moisture availability. We found a high average mean sensitivity value (0.53), which indicated the PAX chronology exhibited enough annual variability to capture fluctuations in environmental conditions. The average interseries correlation (0.74) indicated confident crossdating and a significant association of annual growth among trees within the stand. The positive correlation between the PAX chronology and total precipitation for the local water year was significant (r = 0.53; P < 0.001). Significant positive correlations also were identified between monthly PDSI, monthly total precipitation, and RMJ radial growth. Analyses of temporal stability indicated that the positive relationship between RMJ growth at the PAX site and monthly PDSI was the most stable relationship during the period of analysis (1895–2007). More importantly, we identified a unique inverse relationship between radial growth and monthly mean temperature during periods of the preceding year and current growing year, the first such finding of a strong temperature response for a low-mid elevation tree species in the American Southwest. Our results confirm that RMJ samples collected on the Paxton Springs Malpais are sensitive to climate factors that affect moisture availability, further suggesting that RMJ may be suitable for use in dendroclimatic research at additional locations across the broad distribution of the species.     

An unstable tree-growth response to climate in two 500 year chronologies,North Eastern Qinghai-Tibetan Plateau

Two new Juniper tree-ring-width (TRW) chronologies spanning more than 500 years were developed in the Yellow River source area, North Eastern Qinghai-Tibetan Plateau (NE-QTP). For the two studied sites, located approximately 50 km apart, split correlation and coherence analysis reveal unstable tree-growth responses to local moisture availability. While significant correlations are obtained with April–June local precipitation, Palmer Drought Severity Index (PDSI) and river flow from 1948/1954 to 1998 and from 1948/1954 to 1970s, these correlations vanish for the time period 1970s-1998. The local instrumental climate data (precipitation, PDSI and river flow) exhibit opposite correlations with large scale modes of variability (El Niño Southern Oscillation, ENSO, and Pacific Decadal Oscillation, PDO) before and after the 1977 PDO shift. One tree-ring chronology is coherent and anti-phased with instrumental ENSO/PDO indices at 5.2-year frequency. On the longer time span, this TRW chronology is compared with PDO reconstructed from historical Chinese data. This comparison also exhibits unstable multi-decadal relationships, notably in the mid 19th century. Altogether, the comparison between our two chronologies, local instrumental climate records, and ENSO/PDO indices suggest a cautious use of local TRW records for paleoclimate reconstructions. Further studies are needed to explore both the spatial coherency of tree-ring records and the temporal stability of their response to local and large scale climate variability.     

Different trends and divergent responses to climate factors in the radial growth of Abies georgei along elevations in the central Hengduan Mountains

Climate change has been unprecedented in the last half-century. Tree growth dynamics and responses to climate warming at different elevations vary by study area due to regional diversity in site-specific climatic conditions in the central Hengduan Mountains. A. georgei is the dominant species in high-elevation montane forests in the central Hengduan Mountains. To study the response of A. georgei radial growth to climate and identify tree growth trends at different elevations, tree-ring width chronologies at four elevations across the subalpine A. georgei forest belt were built and growth-climate relationships were analyzed. The primary findings of this study were as follows: (1) radial growth rates of A. georgei decreased with elevation; (2) warming alleviated the limitation of low temperatures and abundant precipitation on tree radial growth at the highest sampling site; and (3) unlike at other elevations, the trend of trees basal area increment (BAI) at the lowest sampling site showed a significant decline over the past 20 years. This suggests the presence of an elevational inflection point, likely between 3800 m and 4000 m, where tree growth trends diverge. These results confirmed that A. georgei at higher elevation in the central Hengduan Mountains currently benefits from higher temperatures. However, the effects of drought on A. georgei at lower elevations would cause radial growth to decrease with climate warming. Therefore, it is critical to establish effective management strategies based on how A. georgei responds to climate change at various elevations.     

Precipitation reconstruction for the southern Altay Mountains (China) from tree rings of Siberian spruce,reveals recent wetting trend

We developed six tree-ring width chronologies of Siberian spruce (Picea obovata) from the low elevation forest of the southern Altay Mountains in northern Xinjiang, China. Although the six chronologies come from different sampling sites, significant correlations existed among the chronologies (r ≥ 0.477), and the first principal component (PC1) accounted for 72.2% of total variance over their common period 1825–2010. Correlation response analysis revealed that radial growth of Siberian spruce is mainly limited by a 12-month precipitation starting from July of the previous year to June of the current year. We therefore developed a July–June precipitation reconstruction spanning 1825–2009, which explained 65.5% of the instrumental variance for the period 1962–2009. The information of our precipitation reconstruction suggested that dry conditions existed for the periods 1829–1838, 1852–1855, 1876–1888, 1898–1911, 1919–1923, 1932–1936, 1943–1955, 1963–1968, 1973–1984 and 2007–2009, and wet conditions for the periods AD 1825–1828, 1839–1851, 1856–1875, 1889–1897, 1912–1918, 1924–1931, 1937–1942, 1956–1962, 1969–1972 and 1985–2006. Spatial climate correlation analyses with gridded land surface data revealed that our precipitation reconstruction contains a strong precipitation signal for the Altay Mountain ranges. Our reconstruction agreed with the moisture-sensitive tree ring width series of Siberian larch from the Altay Mountains of Mongolia on a decadal timescale. In addition, in contrast to a drying trend in north central China, a clear wetting trend has occurred in the southern Altay Mountains since 1980s.     

Leaf growth and population dynamics of intertidal Zostera japonica on the western coast of Korea

Leaf production and population dynamics of Zostera japonica were examined at three elevations of an intertidal transect in Seungbongdo Island on the western coast of Korea. Morphometrics, shoot density, biomass, leaf production, reproductive effort and environmental factors were monitored from October 2001 to October 2002. Z. japonica grew well in the lower intertidal zone from 0.2 to 1.0 m above mean chart datum. The upper station (St. 1) exhibited a finer sediment grain size and richer organic content than the middle (St. 2) and lower stations (St. 3). The size of shoots and leaves was significantly greater at St. 1 than at St. 3, whereas the rhizome internodes were longer at St. 3. Despite differences in morphological characteristics among three stations, seagrass biomass and shoot density were not significantly different among study sites. Shoot density, biomass, morphometrics and leaf productivity exhibited clear seasonal variations, which varied along with seasonal changes of water and air temperature. Leaf productivity measured by the clip and reharvest method was highest in September (4.3 g DW m⁻² d⁻¹) and lowest in February (0.2 g DW m⁻² d⁻¹). Reproductive shoots rapidly increased to maximum density along with the high water temperature in July to September. In the intertidal zone, Z. japonica exhibited faster vegetative growth on muddy sand than on sand, probably due to the difference in nutrient supply. The seasonal changes of water and air temperature were considered to play an important role in the seasonal leaf growth of Z. japonica.     

Intra-annual radial growth of Schrenk spruce (Picea schrenkiana Fisch. et Mey) and its response to climate on the northern slopes of the Tianshan Mountains

Schrenk spruce (Picea schrenkiana Fisch. et Mey.) is widely distributed in the Tianshan Mountains. In this study, four Schrenk spruce trees were continuously monitored with dendrometers from 27 April to 30 September 2014 on the northern slopes of the Tianshan Mountains in northwest China. The goal of this monitoring study was to determine the main growing season of Schrenk spruce and to analyze intra-annual radial growth variability and its relation to daily meteorological factors. Our studies have shown that the critical growing season of Schrenk spruce is from late May to late July and that the rapid growth stage is from mid-June to early July. Meanwhile, in the growing season, changes in the radial growth of Schrenk spruce were negatively correlated with daily temperature, evaporation, sunshine hours and vapor pressure deficit (VPD), and were positively correlated with precipitation and relative humidity (RH). The correlation coefficient between radial growth and RH can be as high as 0.750 (Pearson, p < 0.0001, n = 60). Dates in which precipitation occurred corresponded to periods of rapid growth. The results of the climate-growth analysis show that changes in radial growth reflect the effect of water stress on tree growth, whether or not the changes are positively or negatively correlated with the above climatic factors. This indicates that moisture plays a major role in the growth of Schrenk spruce. We suggest that precipitation between late May to late June is a limiting factor for radial growth of Schrenk spruce on the northern slopes of the Tianshan Mountains.     

Age-related drought sensitivity of Atlas cedar (Cedrus atlantica) in the Moroccan Middle Atlas forests

Age-related tree responses to climate change are still poorly understood at the individual tree level. In this paper, we seek to disentangle the relative contribution of tree age to growth decline and growth–climate relationships in Atlas cedar (Cedrus atlantica Manetti) trees at the Middle Atlas Mountains, northern Morocco. Dendrochronological methods were applied to quantify growth–climate relationships using tree-ring width indices (TRWi) calculated for cedars of two contrasting age groups (old trees, age ≥150 years; young trees, age <150 years). TRWi–climate relationships were assessed at the site and tree levels by using response functions and linear mixed-effects models, respectively. Growth of the studied Atlas cedars was negatively affected by recurrent droughts and by the steep temperature rise since the 1970s. Response functions and mixed-effects models indicated that the decline in tree growth was mainly explained by diminishing precipitation. The negative association between cedar growth and temperature was stronger in old than in young trees. Vulnerability to temperature-induced drought stress in old cedar trees may lead to an impending growth decline. We argue that the age dependence of growth sensitivity to drought must be quantified and considered at the individual tree level when predicting the future dynamics and persistence of cedar forests in the Moroccan Middle Atlas.     

Modeling growth and carbon allocation in two reed beds (Phragmites australis) in the Scheldt estuary

Common reed (Phragmites australis) is a prominent species in the upstream part of the eutrophic Scheldt estuary (Belgium, The Netherlands). From 1996 till 1998, seasonal growth dynamics of the species were studied in two monospecific stands subjected to different salinity regimes (seasonal means 1.6 and 13.3 PSU, respectively). We addressed the following questions: how are these reed vegetations affected by meteorological conditions and by the growth site, what are the important growth processes and what is the fate of the annually fixed carbon. A mathematical model was developed and calibrated using the data from the oligohaline site. Subsequent application of the model to the mesohaline stand required adaptation of parameters relating to the partitioning of resources and timing of growth initiation only. At their peak, the aboveground biomass was 587–1678 g DW m⁻² at the 13.3 PSU site and 1116–2179 g DW m⁻² (1.6 PSU); more than 60% of the biomass was located underground. In 1996, biomasses were 2–3 times lower than in the other 2 years, caused by a retarded growth initiation. Probably due to a lower temperature in early 1996, rhizome bud burst occurred more than 1 month later compared to the other years. In addition, growth initiation was several weeks later in the mesohaline site. This appeared mainly responsible for the large difference in maximal aboveground biomass between both stations. Architecture of the plants was also affected, with a higher shoot density (about 50% more shoots), better-developed root system (15% of total belowground biomass compared to 5%) and more, but smaller leaves at the higher salinity site. Notwithstanding large differences in aboveground biomass, annual growth was similar at both stations (154 and 132 mol C m⁻² per year at the oligo- and mesohaline station, respectively). Primary production accounted for about 80% of all growth processes, rhizome remobilization for almost 20%, translocation of mass before sloughing of leaves accounting about 3%. Within a year, some 44% (oligohaline) and 36% (mesohaline) of new assimilates produced by photosynthesis accumulated as dead litter. The other part was respired by the plant itself, either to provide the energy for growth (23%) or maintenance costs (33–41% at the oligo- and mesohaline station, respectively). Calculated annual turnover rates of aboveground biomass, rhizomes and roots were 100, 62 and 73%, respectively.