Contrasting growth changes in two dominant species of a Mediterranean shrubland submitted to experimental drought and warming

BACKGROUND AND AIMS: Climate projections predict drier and warmer conditions in the Mediterranean basin in the next decades. The possibility of such climatic changes modifying the growth of two Mediterranean species, Erica multiflora and Globularia alypum, which are common components of Mediterranean shrublands, was assessed. METHODS: A field experiment was performed from March 1999 to March 2002 to prolong the drought period and to increase the night-time temperature in a Mediterranean shrubland, where E. multiflora and G. alypum are the dominant species. Annual growth in stem diameter and length of both species was measured and annual stem biomass production was estimated for 1999, 2000 and 2001. Plant seasonal growth was also assessed. KEY RESULTS: On average, drought treatment reduced soil moisture 22 %, and warming increased temperature by 0.7-1.6 degrees C. Erica multiflora plants in the drought treatment showed a 46 % lower annual stem elongation than controls. The decrease in water availability also reduced by 31 % the annual stem diameter increment and by 43 % the annual stem elongation of G. alypum plants. New shoot growth of G. alypum was also strongly reduced. Allometrically estimated biomass production was decreased by drought in both species. Warming treatment produced contrasting effects on the growth patterns of these species. Warmer conditions increased, on average, the stem basal diameter growth of E. multiflora plants by 35 %, raising also their estimated stem biomass production. On the contrary, plants of G. alypum in the warming treatment showed a 14 % lower annual stem growth in basal diameter and shorter new shoots in spring compared with controls. CONCLUSIONS: The results indicate changes in the annual productivity of these Mediterranean shrubs under near future drier and warmer conditions. They also point to alterations in their competitive abilities, which could lead to changes in the species composition of these ecosystems in the long term.     

Warming and drought alter C and N concentration,allocation and accumulation in a Mediterranean shrubland

We investigated the effects of warming and drought on C and N concentrations, nitrogen use efficiency (NUE), and C and N accumulation in different ecosystem compartments. We conducted a 6-year (1999–2005) field experiment to simulate the climate conditions projected by IPCC models for the coming decades in a Mediterranean shrubland. We studied the two dominant species, Globularia alypum and Erica multiflora, and an N-fixing species, Dorycnium pentaphyllum, also abundant in this shrubland. Warming (1 °C) decreased N leaf concentrations by 25% and increased N stem concentrations by 40% in G. alypum. Although warming changed the available ammonium in soil in some seasons, it did not increase total soil N contents. Drought (19% average reduction in soil moisture) decreased leaf N concentrations in the two dominant shrub species, E. multiflora and G. alypum by 16% and 19%, respectively, and increased stem N concentrations by 56% and 40%, respectively. Neither warming nor drought changed the leaf N concentrations in the N-fixing species D. pentaphyllum, although warming increased stem N concentration by 9%. In G. alypum, the increase of stem N concentrations contributed to the observed increase of N accumulation in stem biomass in drought treatments with respect to control plots (8 kg N ha⁻¹). Neither warming nor drought changed NUE in the period 1999–2005. Warming increased soil organic C relative to drought. The effects of warming and drought on C and N concentrations, on N accumulation and on leaf/stem N distribution were not the result of dilution or concentration effects produced by changes in biomass accumulation. Other factors such as the changes in soil N availability, photosynthetic capacity, and plant internal C and N remobilization must be involved. These changes which differed depending on the species and the plant tissue show that the climate change projected for the coming decades will have significant effects on the C and N cycle and stoichiometry, with probable implications for ecosystem structure and function, such as changes in plant–herbivore relationships, decomposition rates or community species composition.     

Precipitation impacts on vegetation spring phenology on the Tibetan Plateau

The ongoing changes in vegetation spring phenology in temperate/cold regions are widely attributed to temperature. However, in arid/semiarid ecosystems, the correlation between spring temperature and phenology is much less clear. We test the hypothesis that precipitation plays an important role in the temperature dependency of phenology in arid/semiarid regions. We therefore investigated the influence of preseason precipitation on satellite‐derived estimates of starting date of vegetation growing season (SOS) across the Tibetan Plateau (TP). We observed two clear patterns linking precipitation to SOS. First, SOS is more sensitive to interannual variations in preseason precipitation in more arid than in wetter areas. Spatially, an increase in long‐term averaged preseason precipitation of 10 mm corresponds to a decrease in the precipitation sensitivity of SOS by about 0.01 day mm^?1. Second, SOS is more sensitive to variations in preseason temperature in wetter than in dryer areas of the plateau. A spatial increase in precipitation of 10 mm corresponds to an increase in temperature sensitivity of SOS of 0.25 day °C^?1 (0.25 day SOS advance per 1 °C temperature increase). Those two patterns indicate both direct and indirect impacts of precipitation on SOS on TP. This study suggests a balance between maximizing benefit from the limiting climatic resource and minimizing the risk imposed by other factors. In wetter areas, the lower risk of drought allows greater temperature sensitivity of SOS to maximize the thermal benefit, which is further supported by the weaker interannual partial correlation between growing degree days and preseason precipitation. In more arid areas, maximizing the benefit of water requires greater sensitivity of SOS to precipitation, with reduced sensitivity to temperature. This study highlights the impacts of precipitation on SOS in a large cold and arid/semiarid region and suggests that influences of water should be included in SOS module of terrestrial ecosystem models for drylands.     

Effects of vegetation canopy and climate on seedling establishment in Mediterranean shrubland

Abstract. Question: Does the influence of plant canopy on seedling establishment interact with climate conditions, and particularly, do intensified drought conditions, enhance a positive effect of the vegetation canopy on seedlings in Mediterranean‐type ecosystems. Location: Mediterranean shrubland near Barcelona, Spain at 210 m a.s.l. Methods: Over the course of four years we recorded seedling emergence and survival in open areas and below vegetation under control, drier and warmer experimental climatic conditions. Results: Seedling emergence is more sensitive to climate conditions than later stages of growth. When considering the whole set of species, the total number of established seedlings at the end of the experiment was lower in the drought and warming stands than in control ones, and vegetation canopy increased the number of these seedlings in the drought stands. Drought reduced seedling emergence but not warming, while the interaction between climate treatments and vegetation canopy was not significant. Seedling survival was lower in the warming treatment than in the control. Under drought conditions, vegetation canopy increased seedling emergence of the dominant Globularia alypum. In control stands, vegetation canopy reduced their survival. Vegetation canopy increased the survival of the dominant Erica multiflora in warming stands, and it reduced the survival of G alypum in drought stands. No significant effects of drought and warming were observed in the seed rain of these two species. Conclusions: The balance of the facilitation‐competition interactions between vegetation canopy and seedling establishment in Mediterranean‐type ecosystems determined by water availability, and drought conditions enhance the positive effect of vegetation canopy. This interaction is species‐specific and shows important between‐year variability.     

The influence of local spring temperature variance on temperature sensitivity of spring phenology

The impact of climate warming on the advancement of plant spring phenology has been heavily investigated over the last decade and there exists great variability among plants in their phenological sensitivity to temperature. However, few studies have explicitly linked phenological sensitivity to local climate variance. Here, we set out to test the hypothesis that the strength of phenological sensitivity declines with increased local spring temperature variance, by synthesizing results across ground observations. We assemble ground‐based long‐term (20–50 years) spring phenology database (PEP725 database) and the corresponding climate dataset. We find a prevalent decline in the strength of phenological sensitivity with increasing local spring temperature variance at the species level from ground observations. It suggests that plants might be less likely to track climatic warming at locations with larger local spring temperature variance. This might be related to the possibility that the frost risk could be higher in a larger local spring temperature variance and plants adapt to avoid this risk by relying more on other cues (e.g., high chill requirements, photoperiod) for spring phenology, thus suppressing phenological responses to spring warming. This study illuminates that local spring temperature variance is an understudied source in the study of phenological sensitivity and highlight the necessity of incorporating this factor to improve the predictability of plant responses to anthropogenic climate change in future studies.     

Drought variation recorded by growth rings of the shrub Sabina vulgaris in the middle Qilian Mountains,northwest China

The evergreen coniferous creeping shrub, Sabina vulgaris, is widely distributed in China, from high and cold mountain sites to low-elevation arid desert areas. This paper performed dendroclimatological analysis of the samples of S. vulgaris from three sampling sites (2700–2900 m a.s.l.) at sites in the northern border of the middle Qilian Mountains, Northwest China. We found that the radial growth of S. vulgaris was mainly limited by rainfall during the growing season, especially in May and June, but was also limited by low temperatures in the late growing season. It presents the regional differention of the climate response of the shrub radial growth from the other distributed areas, such as the sandland and mountains. We constructed a 274-year drought record by calibrating growth-ring data and relating this data to the regional monthly Palmer Drought Severity Index (PDSI₅₆), which represents regional moisture conditions. The PDSI₅₆ reconstruction accounted for 35.5 % of the variation of the actual May to June PDSI₅₆ during the period for which meteorological data was available (1950–2014). The most severe drought occurred during the 1920s, followed by the 1950s, 1740s, and the period from the 1870s to the 1880s; the wettest years were during the 1980s, followed by the 2010s, 1770s, and 1750. The changes between drought, normal, and wet periods were consistent with the reconstructed results from previous conifer ring series for Sabina przewalskii and Picea crassifolia in the study area. Our results demonstrate the value of S. vulgaris in dendrochronology.     

Temperature sensitivity of spring vegetation phenology correlates to within-spring warming speed over the Northern Hemisphere

The inter-annual shift of spring vegetation phenology relative to per unit change of preseason temperature, referred to as temperature sensitivity (days °C⁻¹), quantifies the response of spring phenology to temperature change. Temperature sensitivity was found to differ greatly among vegetation from different environmental conditions. Understanding the large-scale spatial pattern of temperature sensitivity and its underlying determinant will greatly improve our ability to predict spring phenology. In this study, we investigated the temperature sensitivity for natural ecosystems over the North Hemisphere (north of 30°N), based on the vegetation phenological date estimated from NDVI time-series data provided by the Advanced Very High Resolution Radiometer (AVHRR) and the corresponding climate dataset. We found a notable longitudinal change pattern with considerable increases of temperature sensitivity from inlands to most coastal areas and a less obvious latitudinal pattern with larger sensitivity in low latitude area. This general spatial variation in temperature sensitivity is most strongly associated with the within-spring warming speed (WWS; r = 0.35, p < 0.01), a variable describing the increase speed of daily mean temperature during spring within a year, compared with other factors including the mean spring temperature, spring precipitation and mean winter temperature. These findings suggest that the same magnitude of warming will less affect spring vegetation phenology in regions with higher WWS, which might partially reflect plants’ adaption to local climate that prevents plants from frost risk caused by the advance of spring phenology. WWS accounts for the spatial variation in temperature sensitivity and should be taken into account in forecasting spring phenology and in assessing carbon cycle under the projected climate warming.     

北半球中高纬度地区近30年植被春季物候温度敏感性

北半球气候变暖导致植被春季物候开始日期显著提前,温度对春季物候的促进作用是一个过程事件而非瞬时事件,且存在空间差异。该研究在以前研究的基础上,进一步分析温度对植被物候的作用方式,并探讨春季物候温度敏感性的空间特征及影响因素。利用GIMMS3g卫星植被指数产品,采用5种方法提取1982–2009年植被春季物候,并结合格网气象数据计算植被春季物候的温度敏感性,着重分析自然植被春季物候温度敏感性与环境因素的关系。结果表明,温度是北半球植被春季物候的主要制约因素,54%的像元显示温度最大效应发生在物候开始当月和之前一个月。温度主导的春季物候的像元中,91.3%的像元指示早春温度对物候开始的促进作用。植被春季物候的温度敏感性存在空间异质性,随着区域环境因素的不同,年际温度标准差、累积降水量和辐射对植被春季物候温度敏感性都具有各自或协同的调控作用。     

Climate-induced long-term changes in the phenology of Mediterranean fungi

Long-term responses in the phenology of Mediterranean macrofungi to climatic changes are poorly documented. Here, we address this issue by comparing the fruiting patterns of 159 fungal species in Southern France between the first half of the 19th century and the first decade of the 21st century. We used a trait-based approach to assess the influence of phenology and morphology of fungal fruit bodies and their site ecology and biogeography on the response to climate change. We show that early autumnal fruiters, epigeous species and species with affinities for cold climates now start to fruit on average 16.4, 17.3 and 17.3 d later compared to their emergence dates in the 19th century, while late fruiters, hypogeous species and Mediterranean-restricted species did not change their fruiting date. Among ecological guilds, saproxylic species and pine-associated mutualists delayed their autumnal emergence by 32.5 and 19.2 d, likely in response to a delayed rewetting of litter and woody debris after extended summer drought. Our results suggest that long-term climate warming in the Mediterranean was accompanied by contrasting changes in the emergence of fungal fruit bodies according to ecological guilds, sporocarp life-forms and forest types.     

Changes in the onset of spring growth in shrubland species in response to experimental warming along a north–south gradient in Europe

Aim To test whether the onset of spring growth in European shrublands is advanced in response to the warmer conditions projected for the next two decades by climate models, and, if there is a change, whether it differs across Europe. Location The studied sites spanned a broad north–south European gradient with average annual temperatures (8.2–15.6 °C) and precipitation (511–1427 mm). Methods ‘Bud break’ was monitored in eight shrub and grass species in six European sites under control and experimentally warmer conditions generated by automatic roofs covering vegetation during the night. Results Species responsive to increased temperatures were Vaccinium myrtillus and Empetrum nigrum in Wales, Deschampsia flexuosa in Denmark, Calluna vulgaris in Netherlands, Populus alba in Hungary and Erica multiflora in Spain. Although the acceleration of spring growth was the commonest response to warming treatments, the responses at each site were species specific and year dependent. Under experimental warming 25% of cases exhibited a significantly earlier onset of the growing season and 10% had a significantly delayed onset of vegetative growth. No geographical gradient was detected in the experimental warming effects. However, there was a trend towards a greater dominance of phenological advances with more intense the warming treatments. Above 0.8 °C warming, only advancements were recorded. Main conclusions Our results show that warmer temperatures projected for the next decades have substantial potential effects on the phenology of the spring growth of dominant species in different European shrublands, with a dominant trend towards advancements the more intense the warming is. However, our study also demonstrates the overall difficulties of applying simple predictive relationships to extrapolate the effects of global change on phenology. Various combinations of environmental factors occur concurrently at different European sites and the interactions between different drivers (e.g. water and chilling) can alter phenology significantly.     

Delayed autumn phenology in the Northern Hemisphere is related to change in both climate and spring phenology

The timing of the end of the vegetation growing season (EOS) plays a key role in terrestrial ecosystem carbon and nutrient cycles. Autumn phenology is, however, still poorly understood, and previous studies generally focused on few species or were very limited in scale. In this study, we applied four methods to extract EOS dates from NDVI records between 1982 and 2011 for the Northern Hemisphere, and determined the temporal correlations between EOS and environmental factors (i.e., temperature, precipitation and insolation), as well as the correlation between spring and autumn phenology, using partial correlation analyses. Overall, we observed a trend toward later EOS in ~70% of the pixels in Northern Hemisphere, with a mean rate of 0.18 ± 0.38 days yr^?1. Warming preseason temperature was positively associated with the rate of EOS in most of our study area, except for arid/semi‐arid regions, where the precipitation sum played a dominant positive role. Interestingly, increased preseason insolation sum might also lead to a later date of EOS. In addition to the climatic effects on EOS, we found an influence of spring vegetation green‐up dates on EOS, albeit biome dependent. Our study, therefore, suggests that both environmental factors and spring phenology should be included in the modeling of EOS to improve the predictions of autumn phenology as well as our understanding of the global carbon and nutrient balances.     

Climate change and shifts in spring phenology of three horticultural woody perennials in northeastern USA

We evaluated spring phenology changes from 1965 to 2001 in northeastern USA utilizing a unique data set from 72 locations with genetically identical lilac plants (Syringa chinensis, clone Red Rothomagensis). We also utilized a previously validated lilac-honeysuckle spring index model to reconstruct a more complete record of first leaf date (FLD) and first flower date (FFD) for the region from historical weather data. In addition, we examined mid-bloom dates for apple (Malus domestica) and grape (Vitis vinifera) collected at several sites in the region during approximately the same time period. Almost all lilac sites with significant linear trends for FLD or FFD versus year had negative slopes (advanced development). Regression analysis of pooled data for the 72 sites indicated an advance of –0.092 day/year for FFD (P=0.003). The slope for FLD was also negative (–0.048 day/year), but not significant (P=0.234). The simulated data from the spring index model, which relies on local daily temperature records, indicated highly significant (P<0.001) negative slopes of –0.210 and –0.123 day/year for FLD and FFD, respectively. Data collected for apple and grape also indicated advance spring development, with slopes for mid-bloom date versus year of –0.20 day/year (P=0.01) and –0.146 (P=0.14), respectively. Collectively, these results indicate an advance in spring phenology ranging from 2 to 8 days for these woody perennials in northeastern USA for the period 1965 to 2001, qualitatively consistent with a warming trend, and consistent with phenology shifts reported for other mid- and high-latitude regions.     

Water status and growth phenology of a Saharan shrub in north Africa

An experimental study of the relationship between the phenological phases, water relations and climatic conditions was conducted on one shrub species, Spartidium saharae, in Saharan ecosystem of southern Tunisia. The results show that the water potential remained significantly higher than that recorded in other desert shrubs. However, a relationship between the water potential of S. saharae and precipitation was not evident suggesting that access to permanent ground water may be a factor of this decoupling. Although shoot elongation of the species occurred outside the wet season, water shortage induced a decrease in fruit production.     

黄花倒水莲生长与内源激素水平动态变化关系研究北大核心CSCD

为探索黄花倒水莲春梢生理生化特性的差异以及不同内源激素的变化规律,该文对黄花倒水莲春梢的生长动态进行监测,采用间接酶联免疫吸附法(ELISA)测定脱落酸(ABA)、生长素(IAA)、赤霉素(GA)、乙烯(ETH)和玉米素核苷(ZR)五种内源激素含量的动态变化,并对两者间的相关性进行分析。结果表明:(1)黄花倒水莲春梢生长发育过程可分为快速增长期(0~12 d)、生长转折期(16~20 d)和缓慢增长期(24~32 d)三个阶段。(2)内源激素ABA、GA、ETH和ZR含量在缓慢增长期显著高于快速增长期和生长转折期,IAA含量各时期差异较小。(3)春梢长、底部叶长和叶宽在快速增长期与ABA、GA、ETH和ZR含量呈负相关,且与ZR含量具有一定显著性,与IAA含量呈正相关;生长转折期,各指标与GA、ETH和ZR含量呈正相关,与GA含量具有一定显著性,与ABA含量呈负相关;缓慢增长期,各指标与五种内源激素含量均呈正相关,与IAA和ZR含量具有一定显著性。该研究结果为生产上利用外源激素调控黄花倒水莲春梢抽出以及生长提供了理论基础。     

Leaf area and crop growth in relation to phenology of barley in Mediterranean environments

In a barley/livestock farming system of northern Syria, high biomass production in addition to high grain yield is desirable. The aim of this study was to assess the effect of environment and phenology on growth and yield of barley in northern Syria. Leaf area duration (LAD), specific leaf area (SLA), crop growth rate (CGR) on a thermal time basis, and biological and grain yields were compared for entries representing three contrasting development patterns: early heading spring types (pattern A), medium early heading winter types (pattern B), and late heading spring types (pattern C). The experiment was conducted in 1988/89 at two sites: Tel Hadya (TH, 250 mm precipitation) and Breda (BR, 180 mm). Cold damage occurred in winter and, especially at BR, drought stress developed in spring.At the two sites, development was similar, but yields at TH were twice those at BR. This was related to a longer LAD and a faster CGR in spring. Development pattern affected growth. A long vegetative phase (pattern B) resulted in small leaves with a low SLA in winter, probably due to a slow leaf extension rate. Since cold tolerance and profuse tillering compensated for the small leaf size, pattern B had on average a longer LAD than pattern A. Pattern C had a longer LAD than pattern A because of a longer crop duration. This long duration had a negative effect on yield, so LAD was poorly related to yield. Development in spring was associated with CGR. Pattern C had a slow CGR and low yields; pattern B had the fastest CGR, but the yield advantage over pattern A was not significant. These results suggest that early heading winter barley, which combines long LAD with fast spring CGR, may give the best performance in a barley/livestock farming system in northern Syria.     

The importance of phenology for the evaluation of impact of climate change on growth of boreal, temperate and Mediterranean forests ecosystems: an overview

An overview is presented of the phenological models relevant for boreal coniferous, temperate-zone deciduous and Mediterranean coniferous forest ecosystems. The phenology of the boreal forests is mainly driven by temperature, affecting the timing of the start of the growing season and thereby its duration, and the level of frost hardiness and thereby the reduction of foliage area and photosynthetic capacity by severe frost events. The phenology of temperate-zone forests is also mainly driven by temperature. Since temperate-zone forests are mostly mixed-species deciduous forests, differences in phenological response may affect competition between tree species. The phenology of Mediterranean coniferous forests is mainly driven by water availability, affecting the development of leaf area, rather than the timing of phenological events. These phenological models were subsequently coupled to the process-based forest model FORGRO to evaluate the effect of different climate change scenarios on growth. The results indicate that the phenology of each of the forest types significantly affects the growth response to a given climate change scenario. The absolute responses presented in this study should, however, be used with caution as there are still uncertainties in the phenological models, the growth models, the parameter values obtained and the climate change scenarios used. Future research should attempt to reduce these uncertainties. It is recommended that phenological models that describe the mechanisms by which seasonality in climatic drivers affects the phenological aspects of trees should be developed and carefully tested. Only by using such models may we make an assessment of the impact of climate change on the functioning and productivity of different forest ecosystems. Received: 21 October 1999 / Revised: 10 May 2000 / Accepted: 10 May 2000     

1982—2009年基于卫星数据的北半球中高纬地区植被春季物候动态及其与气候的关系

深入认识北半球植被物候在全球变暖背景下的动态变化特征,对于评估和预测生态系统结构和功能对气候变化的响应有重要的指示作用.遥感技术是获取北半球植被春季物候的最重要方法,但是由于物候提取算法的差异,目前还存在较大的不确定性.本文利用5种方法,基于卫星获取的归一化植被指数估算了北半球中高纬地区1982—2009年植被春季物候开始日期,分析了该日期的多年动态变化的时空特征,并探讨了气候变化对春季物候变化的影响.结果表明: 研究区植被春季物候开始日期呈现提前趋势,研究期间提前(4.0±0.8) d,其中,欧亚大陆提前速率为(0.22±0.04) d·a^-1,显著高于北美大陆的变化速率(0.03±0.02 d·a^-1);不同植被类型的变化趋势不同,5种方法都显示草地表现为显著提前趋势,而林地的提前趋势不显著.区域平均的植被春季物候开始日期的年际波动主要受春季温度的变化所驱动(r² =0.61,P<0.001), 温度每上升1 ℃,可以导致春季物候提前(3.2±0.5) d,而春季降水影响不显著(P>0.05).     

Climate change and timing of spring migration in the long-distance migrant Ficedula hypoleuca in central Europe: the role of spatially different temperature changes along migration routes

Coinciding with increasing spring temperatures in Europe, many migrants have advanced their arrival or passage times over the last decades. However, some species, namely long-distance migrants, could be constrained in their arrival dates due to their largely inherited migratory behaviour and thus a likely inflexibility in their response to exogenous factors. To examine this hypothesis for pied flycatchers (Ficedula hypoleuca), we tested the effects of the temperature regimes along their migration routes north of the Sahara on their arrival times in central Europe. To do so, we developed a site-independent large-scale approach based on temperature data available on the Internet. Temperature regimes along the migration routes of pied flycatchers within Europe convincingly correlate with their first arrival times. It can be concluded that the progression of spring migration in this species is strongly influenced by temperature en route. Because of the recent inconsistent climatic changes in various parts of Europe, we hypothesize that individuals migrating along different routes will be unequally affected by further climatic changes.