Environmental constraints on phenology and internal nutrient cycling in the Mediterranean winter-deciduous shrub Amelanchier ovalis Medicus

The functional adjustments of winter-deciduous perennials to Mediterranean conditions have received little attention. The objectives of this study were: (i) to determine whether Amelanchier ovalis, a winter-deciduous shrub of Mediterranean and sub-Mediterranean regions, has nutritional and phenological traits in common with temperate zone deciduous trees and shrubs and (ii) to determine the constraints of Mediterranean environmental conditions on these traits. Over two years, phenology and nitrogen, and phosphorus concentrations were monitored monthly in the crown of A. ovalis. Leaf longevity, survival and nutrient resorption from senescing leaves were used to infer nutrient use efficiency and retention times of nutrients within the crown. In A. ovalis, bud burst was much earlier than in temperate deciduous trees and shrubs. Most vegetative and reproductive growth occurred in spring. Limited phenological development took place during the summer drought period. Unexpectedly, leaf shedding was very gradual, which might be related to water shortages in summer. Leaf longevity, nutrient resorption from senescing leaves, and maximum leaf nutrient concentrations indicated that nutrient retention times were short and nutrient use efficiency was low compared to that found in temperate deciduous plants and co-occurring Mediterranean evergreens. A. ovalis exhibited phenological development appropriate for a Mediterranean climate, although its limited ability to retain nutrients likely restricts the types of sites that it can occupy.     

Seasonal branch nutrient dynamics in two Mediterranean woody shrubs with contrasted phenology

BACKGROUND AND AIMS: Mediterranean woody plants have a wide variety of phenological strategies. Some authors have classified the Mediterranean phanaerophytes into two broad phenological categories: phenophase-overlappers (that overlap resource-demanding activities in a short period of the year) and phenophase-sequencers (that protract resource-demanding activities throughout the year). In this work the impact of both phenological strategies on leaf nutrient accumulation and retranslocation dynamics at the level of leaves and branches was evaluated. Phenophase-overlappers were expected to accumulate nutrients in leaves throughout most of the year and withdraw them efficiently in a short period. Phenophase-sequencers were expected to withdraw nutrients progressively throughout the year, without long accumulation periods. METHODS: To test this hypothesis, variations in phenology and leaf NPK in the crown of a phenophase-overlapper Cistus laurifolius and a phenophase-sequencer Bupleurum fruticosum were monitored monthly during 2 years. KEY RESULTS: Changes in nutrient concentration at the leaf level were not clearly related with the different phenologies. Nitrogen and phosphorous resorption efficiencies were lower in the phenophase-overlapper, and accumulation-retranslocation seasonality was similar in both species. Changes in the branch nutrient pool agreed with the hypothesis that the phenophase-overlapper accumulated nutrients from summer until the bud burst of the following spring, recovering a large nutrient pool during massive leaf shedding. The phenophase-sequencer did not accumulate nutrients from autumn until early spring, achieving lower nutrient recovery during spring leaf shedding. CONCLUSIONS: It is concluded that phenological demands influence branch nutrient cycling. This effect is easier to detect by assessing changes in the branch nutrient pool rather than changes in the leaf nutrient concentration.     

Influence of land-use types and climatic variables on seasonal patterns of NDVI in Mediterranean Iberian ecosystems

Question: What is the influence of management on the functioning of vegetation over time in Mediterranean ecosystems under different climate conditions? Location: Mediterranean shrublands and forests in SE Iberia (Andalusia). Methods: We evaluated the Normalized Difference Vegetation Index (NDVI) for the 1997-2002 time series to determine phenological vegetation patterns under different historical management regimes. Three altitudinal ranges were considered within each area to explore climate × management interactions. Each phenological pattern was analysed using time series statistics, together with precipitation (monthly and cumulative) and temperature. Results: NDVI time series were significantly different under different management regimes, particularly in highly transformed areas, which showed the lowest NDVI, weakest annual seasonality and a more immediate phenological response to precipitation. The NDVI relationship with precipitation was strongest in the summer-autumn period, when precipitation is the main plant growth-limiting factor. Conclusions: NDVI time series analyses elucidated complex influences of land use and climate on ecosystem functioning in these Mediterranean ecosystems. We demonstrated that NDVI time series analyses are a useful tool for monitoring programmes because of their sensitivity to changes, ease of use and applicability to large-scale studies.     

Asphodelus aestivus,an example of synchronization with the climate periodicity

The phenology ofAsphodelus aestivus Brot. is described by means of a phenological model which has been formulated to fit skewed phenological data. Based on the model parameters the timing of different phenophases of biomass accumulation were determined. The biomass oscillations of leaves, inflorescence stalks and tubers were found to be synchronized with the predictable seasonal climatic changes. In addition, the plant seems to respond to minor random climatic variations. The emergence of leaves and inflorescence stalks depends on stored material in the tubers while leaf and inflorescence stalk elongation as well as flowering depends on current production. The storage part of the tubers seems to be a regulating structure, which is responsible for the synchronization ofA. aestivus productivity with the seasonality of the Mediterranean climate.A. aestivus is considered to be a Competitor Ruderal and Stress tolerant (C-R-S) strategist which may explain the wide distribution of this plant over the Mediterranean Basin.     

植物物候学研究进展

植物物候变化在研究陆地生态系统对气候变化的响应时被誉为"矿井中的金丝雀",全球气候变化愈演愈烈,重新引起了人们对植物物候研究的广泛关注。随着观测技术的发展,在各种空间和生态尺度上收集到的物候观测数据迅速累积,尽管已经在多个尺度上（物种、群落和景观尺度）观察到物候变化,但物候变化的机理仍然没有得到很好的理解。回顾了国内外植物物候研究的发展历程;总结了物候数据收集技术进展和全球物候变化的主要趋势;归纳了植物物候变化的机理与驱动因素;探讨了物候模型研究及物候对气候变化响应研究的主要方向。随着物候观测技术在不同尺度上应用的增加,物候研究进入了一个新的阶段。未来物候研究需要制定跨区域标准化观测指南,融合所有相关学科,改进物候模型,拓展研究区域;同时融合有效的历史物候资料,采用新技术和长期收集的物候数据为大数据时代植物物候学研究提供基础。     

Variations in the timing of different phenological stages of cambial activity in Abies pindrow (Royle) along an elevation gradient in the north-western Himalaya

High mountains around the globe are some of the most vulnerable ecosystems to climate change and of great concern for conservation. The Himalayan Mountains are experiencing a higher warming than average global warming, which can significantly impact their biodiversity, vegetation distribution and ecosystem structure. There is a need to study the process of wood formation in Himalayan conifers to have a better understanding of their growth responses to predicted climate change. Variations in the climatic factors regulating cambial activity would result in changes in the timing of cambial phenology. In this study, the variations in the timing of different stages of cambial phenology (cell enlargement stage, wall-thickening stage and cell maturation stages) in pindrow fir (Abies pindrow) were investigated from anatomical observations of wood microcores collected during 2014-15 along an elevation range of c.2300−3000 m asl in the north-western Himalaya. The onset of all three cambial phenological stages was significantly correlated with elevation, with onset of cambial activity happening more than a week earlier at the lowest elevation than at the highest elevation site. Although the termination of wall-thickening and maturation stage appeared minimally related to elevation, the cell-enlargement stage showed significant correlation with elevation, with tracheid formation ceasing approximately three weeks earlier in trees at the highest elevation. The timing of these phenological stages did not show strong variations between the two study years. Our findings provide new data on the timings of cambial phenophases and help to understand tree growth response to ongoing changing climate in the Himalayan region.     

Impact of climate change on plant phenology in Mediterranean ecosystems

Plant phenology is strongly controlled by climate and has consequently become one of the most reliable bioindicators of ongoing climate change. We used a dataset of more than 200 000 records for six phenological events of 29 perennial plant species monitored from 1943 to 2003 for a comprehensive assessment of plant phenological responses to climate change in the Mediterranean region. Temperature, precipitation and North Atlantic Oscillation (NAO) were studied together during a complete annual cycle before phenological events to determine their relative importance and potential seasonal carry‐over effects. Warm and dry springs under a positive phase of NAO advance flowering, leaf unfolding and fruiting dates and lengthen the growing season. Spatial variability of dates (range among sites) was also reduced during warm and dry years, especially for spring events. Climate during previous weeks to phenophases occurrence had the greatest impact on plants, although all events were also affected by climate conditions several months before. Immediate along with delayed climate effects suggest dual triggers in plant phenology. Climatic models accounted for more than 80% of variability in flowering and leaf unfolding dates, and in length of the growing season, but for lower proportions in fruiting and leaf falling. Most part of year‐to‐year changes in dates was accounted for temperature, while precipitation and NAO accounted for <10% of dates' variability. In the case of flowering, insect‐pollinated species were better modelled by climate than wind‐pollinated species. Differences in temporal responses of plant phenology to recent climate change are due to differences in the sensitivity to climate among events and species. Spring events are changing more than autumn events as they are more sensitive to climate and are also undergoing the greatest alterations of climate relative to other seasons. In conclusion, climate change has shifted plant phenology in the Mediterranean region.     

Climate changes affecting biotic interactions,phenology, and reproductive success in a savanna community over a 10-year period

Climatic parameters are able to influence the timing of phenological events affecting the degree of synchrony among plant species, their interactions, and reproductive success. Shrubs of Malpighiaceae family in the Brazilian Tropical Savanna present sequential flowering phenology. We verified variations in climatic factors (temperature and precipitation) over a period of 10 years (2005–2014) and correlated them with the onset of flowering of four of these Malpighiaceae species. Furthermore, we tested whether the phenological synchronization among species has changed over time affecting the herbivory and fruit set. Herbivory and fruit production were recorded during three reproductive seasons (2008/2009, 2011/2012, 2013/2014). We developed a mathematical model to estimate the flower and fruit production in response to phenological changes for the next 5 years. Results show that climatic factors changed, influencing the onset of species flowering. The degree of overlap among species also changed and the effects on species interactions were species specific. The mathematical model successfully presented a tendency on flower and fruit production contributing to the predictions of the outcomes in response to phenological changes. We confirm the effects of climate changes on plant phenological events and the importance of feature plasticity for better performance of species.     

日光诱导叶绿素荧光对亚热带常绿针叶林物候的追踪

植被物候期(春季返青和秋季衰老)是表征生物响应和陆地碳循环的基础信息。由于常绿针叶林冠层绿度的季节变动较弱,遥感提取常绿针叶林的物候信息存在着较大的不确定性,是目前区域物候监测中的难点。利用MODIS植被指数(归一化植被指数NDVI和增强型植被指数EVI)、GOME-2日光诱导叶绿素荧光(SIF)和通量数据(总初级生产力GPP)估算2007—2011年亚热带常绿针叶林物候期,用来比较三类遥感指数估算常绿针叶林物候的差异。结果表明:基于表征光合作用物候的通量GPP数据估算得到5年内亚热带常绿针叶林生长季开始时间(SOS_GPP)为第63天,生长季结束时间(EOS_GPP)为第324天,生长季长度为272天;基于反映植被光合作用特征的SIF曲线获得物候信息要滞后GPP物候期,其中生长季开始时间滞后19天,生长季结束时间滞后2天;基于传统植被指数NDVI和EVI的物候期滞后GPP物候期的时间要大于SIF滞后期,其中植被指数SOS滞后SOS_GPP31天,植被指数EOS滞后EOS_GPP10—17天。虽然基于3种遥...     

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     

Genetic diversity increases regional variation in phenological dates in response to climate change

Climate change is inducing changes in the phenological timings of organisms. Genetic diversity could influence phenological responses to climate change, but empirical evidence is very limited. We estimated the regional variation across Japan in flowering and leaf budburst dates of plants based on a dataset of phenological timings from 1953 to 2005. The observed plants' genetic diversities varied according to human cultivation. The within-species variations of phenological response to temperature as well as regional variations were less in the plant populations with lower genetic diversity. Thus, genetic diversity influences the variation in phenological responses of plant populations. Under increased temperatures, low variation in phenological responses may allow drastic changes in the phenology of plant populations with synchronized phenological timings. Our findings indicate that we should pay attention to maintaining genetic diversity of populations to alleviate changes in phenology due to future climate change.     

Influence of Habitat and Climate Variables on Arbuscular Mycorrhizal Fungus Community Distribution,as Revealed by a Case Study of Facultative Plant Epiphytism under Semiarid Conditions

In semiarid Mediterranean ecosystems, epiphytic plant species are practically absent, and only some species of palm trees can support epiphytes growing in their lower crown area, such as Phoenix dactylifera L. (date palm). In this study, we focused on Sonchus tenerrimus L. plants growing as facultative epiphytes in P. dactylifera and its terrestrial forms growing in adjacent soils. Our aim was to determine the possible presence of arbuscular mycorrhizal fungi (AMF) in these peculiar habitats and to relate AMF communities with climatic variations. We investigated the AMF community composition of epiphytic and terrestrial S. tenerrimus plants along a temperature and precipitation gradient across 12 localities. Epiphytic roots were colonized by AMF, as determined by microscopic observation; all of the epiphytic and terrestrial samples analyzed showed AMF sequences from taxa belonging to the phylum Glomeromycota, which were grouped in 30 AMF operational taxonomic units. The AMF community composition was clearly different between epiphytic and terrestrial root samples, and this could be attributable to dispersal constraints and/or the contrasting environmental and ecophysiological conditions prevailing in each habitat. Across sites, the richness and diversity of terrestrial AMF communities was positively correlated with rainfall amount during the most recent growing season. In contrast, there was no significant correlation between climate variables and AMF richness and diversity for epiphytic AMF communities, which suggests that the composition of AMF communities in epiphytic habitats appears to be largely determined by the availability and dispersion of fungal propagules from adjacent terrestrial habitats.     

Phenological responses of plants to climate change in an urban environment

Global climate change is likely to alter the phenological patterns of plants due to the controlling effects of climate on plant ontogeny, especially in an urbanized environment. We studied relationships between various phenophases (i.e., seasonal biological events) and interannual variations of air temperature in three woody plant species (Prunus davidiana, Hibiscus syriacus, and Cercis chinensis) in the Beijing Metropolis, China, based on phenological data for the period 1962–2004 and meteorological data for the period 1951–2004. Analysis of phenology and climate data indicated significant changes in spring and autumn phenophases and temperatures. Changes in phenophases were observed for all the three species, consistent with patterns of rising air temperatures in the Beijing Metropolis. The changing phenology in the three plant species was reflected mainly as advances of the spring phenophases and delays in the autumn phenophases, but with strong variations among species and phenophases in response to different temperature indices. Most phenophases (both spring and autumn phenophases) had significant relationships with temperatures of the preceding months. There existed large inter- and intra-specific variations, however, in the responses of phenology to climate change. It is clear that the urban heat island effect from 1978 onwards is a dominant cause of the observed phenological changes. Differences in phenological responses to climate change may cause uncertain ecological consequences, with implications for ecosystem stability and function in urban environments.     

Trophic level asynchrony in rates of phenological change for marine,freshwater and terrestrial environments

Recent changes in the seasonal timing (phenology) of familiar biological events have been one of the most conspicuous signs of climate change. However, the lack of a standardized approach to analysing change has hampered assessment of consistency in such changes among different taxa and trophic levels and across freshwater, terrestrial and marine environments. We present a standardized assessment of 25 532 rates of phenological change for 726 UK terrestrial, freshwater and marine taxa. The majority of spring and summer events have advanced, and more rapidly than previously documented. Such consistency is indicative of shared large scale drivers. Furthermore, average rates of change have accelerated in a way that is consistent with observed warming trends. Less coherent patterns in some groups of organisms point to the agency of more local scale processes and multiple drivers. For the first time we show a broad scale signal of differential phenological change among trophic levels; across environments advances in timing were slowest for secondary consumers, thus heightening the potential risk of temporal mismatch in key trophic interactions. If current patterns and rates of phenological change are indicative of future trends, future climate warming may exacerbate trophic mismatching, further disrupting the functioning, persistence and resilience of many ecosystems and having a major impact on ecosystem services.     

Vulnerability to forest loss through altered postfire recovery dynamics in a warming climate in the Klamath Mountains

In the context of ongoing climatic warming, certain landscapes could be near a tipping point where relatively small changes to their fire regimes or their postfire forest recovery dynamics could bring about extensive forest loss, with associated effects on biodiversity and carbon‐cycle feedbacks to climate change. Such concerns are particularly valid in the Klamath Region of northern California and southwestern Oregon, where severe fire initially converts montane conifer forests to systems dominated by broadleaf trees and shrubs. Conifers eventually overtop the competing vegetation, but until they do, these systems could be perpetuated by a cycle of reburning. To assess the vulnerability of conifer forests to increased fire activity and altered forest recovery dynamics in a warmer, drier climate, we characterized vegetation dynamics following severe fire in nine fire years over the last three decades across the climatic aridity gradient of montane conifer forests. Postfire conifer recruitment was limited to a narrow window, with 89% of recruitment in the first 4 years, and height growth tended to decrease as the lag between the fire year and the recruitment year increased. Growth reductions at longer lags were more pronounced at drier sites, where conifers comprised a smaller portion of live woody biomass. An interaction between seed‐source availability and climatic aridity drove substantial variation in the density of regenerating conifers. With increasing climatic water deficit, higher propagule pressure (i.e., smaller patch sizes for high‐severity fire) was needed to support a given conifer seedling density, which implies that projected future increases in aridity could limit postfire regeneration across a growing portion of the landscape. Under a more severe prospective warming scenario, by the end of the century more than half of the area currently capable of supporting montane conifer forest could become subject to minimal conifer regeneration in even moderate‐sized (10s of ha) high‐severity patches.     

Restoring biodiversity to pine afforestations in Israel

This paper presents a practitioner's perspective of the Keren Kayemeth Leisrael's (KKL – Israel's Forest Service) forest management policy, including actions encouraging biological diversity attributes to Israel's planted conifer forests. These changes are reviewed in light of institutional changes within the KKL, recent global initiatives and scientific trends concerning biodiversity, ecologically oriented forestry and sustainable forest management.

The management of Israel's planted conifer forests for biological diversity values is a relatively new phenomenon. Most of Israel's high forests were planted and consist primarily of a small core group of native and exotic Mediterranean conifers. Over time, these simplified afforestations evolved into a complex set of forest stands – a “near-native” type of forest ecosystem embodying a sum total of natural and artificial processes. They can thus serve as models to help visualise and understand how plantation-type forests can be converted into complex afforestations systems possessing a higher degree of structural, functional, compositional and genetic diversity.     

Mediterranean climate effects. I. Conifer water use across a Sierra Nevada ecotone

Xylem water potential of the midelevation conifers Pinus jeffreyi, Pinus lambertiana, Abies concolor, and Calocedrus decurrens, the higher elevation Pinus monticola and Abies magnifica, and co-occurring evergreen angiosperm shrubs, together with soil moisture under these plants, were monitored at three sites on the Kern Plateau in the southernmost Sierra Nevada Range of California. Site locations spanned the ecotone between the mid- and upper montane forests at elevations of 2230-2820 m. Measurements were made through a low-snowfall year and a heavy-snowfall year.In the Mediterranean climate of the Sierra Nevada, the heavy winter snowpack persists into late spring, after precipitation has effectively stopped. We found the subsequent depletion of soil moisture due to plant water uptake to result in predawn xylem water potentials for conifers more negative by 0.6-1.4 MPa than those for shrubs or inferred soil potentials. Shrubs generally depleted soil moisture more rapidly and ultimately extracted a greater fraction of the available soil moisture than did the conifers. This depletion of soil moisture by shrubs, particularly Arctostaphylos patula, may limit conifer growth and regeneration by prematurely terminating growth on the shallow soils studied. The conifers all generally showed similar patterns of soil moisture use, except that A. magnifica extracted moisture more rapidly early in the season.     

Arboreal and prostrate conifers coexisting in Mediterranean high mountains differ in their climatic responses

In contrast to most high elevation areas, plant growth at Mediterranean mountains is exposed to a summer drought period, which represents an additional climatic constraint to low temperatures. Although arboreal and shrubby conifers coexist at high altitudes, most dendroecological studies have focused on climatic responses of tree species, whereas those of shrubby species remain mostly unexplored. We built tree-ring width chronologies for two conifer species, a shrub (Juniperus sabina) and a tree (Pinus sylvestris), coexisting at three high-altitude localities of the Iberian System mountains, eastern Spain. We analyzed their climate–growth relationships for the period 1950–2009 using correlation analyses and multiple regressions. Coexisting species responded to year-to-year climatic variability in different ways. Radial growth in junipers and pines responded positively to April and May temperatures, respectively. Summer drought constrained growth in both cases, although its impact was stronger on junipers than on pines. Our findings suggest that junipers respond earlier than pines to spring temperatures due to their prostrate morphology which may enhance a fast warming of their cambial meristems after snowmelt. The higher dependence of J. sabina on summer rainfall as compared with co-occurring pines confirms that drought stress negatively impacts secondary growth in Mediterranean mountains. This sensitivity to water availability may be caused by the juniper shallow root systems, which mainly use superficial soil water. The climatic signal registered in J. sabina allows studying the response of other similar shrubby woody species growing in Mediterranean alpine areas to the ongoing climate warming, which could also reduce water availability.     

Deriving a new phenological indicator of interannual net carbon exchange in contrasting boreal deciduous and evergreen forests

Phenology is an important variable affecting the annual net ecosystem production (NEP) of terrestrial ecosystems. A new phenological indicator was proposed based on the ratio of respiration season length and growing season length (respiration–growth length ratio, RGR). Validation of this new phenological indicator was conducted using continuous flux measurements at contrasting boreal deciduous and evergreen forests in Canada. Analyses based on yearly anomalies of both annual NEP and phenological indicators indicated that the RGR can explain more proportion of interannual NEP variability compared to existing phenological metrics, including the carbon uptake period and the autumn lag. A multivariate regression model was used to predict the respiration–growth length ratio anomaly using anomalies of spring air temperature, autumn radiation and soil water content (SWC), which serves as a prerequisite for this indicator being scaled up for regional applications where flux data were unavailable. By normalization growing season length, interannual NEP showed comparable sensitivity to RGR variations of different plant functional types, which is a great advantage over other phenological indicators. The high potential of RGR in explaining interannual NEP variability may highlight the importance of respiration process in controlling annual NEP, which has probably been overlooked or underestimated in existing phenological studies. The comparable sensitivity of RGR to annual NEP observed at different plant functional types would favor its application in tracking interannual variability of NEP regionally and complementary to existing indices to promote our understanding of carbon sequestration with future climate change.