Article

Ecological Impacts of a Widespread Frost Event Following Early Spring Leaf-Out

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Abstract

In the spring of 2010, temperatures averaged ~3 °C above the long-term mean (March–May) across the northeastern United States. However, in mid-to-late spring, much of this region experienced a severe frost event. The spring of 2010 therefore provides a case study on how future spring temperature extremes may affect northeastern forest ecosystems. We assessed the response of three northern hardwood tree species (sugar maple, American beech, yellow birch) to these anomalous temperature patterns using several different data sources and addressed four main questions: (1) Along an elevational gradient, how was each species affected by the late spring frost? (2) How did differences in phenological growth strategy influence their response? (3) How did the late spring frost affect ecosystem productivity within the study domain? (4) What are the potential long-term impacts of spring frost events on forest community ecology? Our results show that all species exhibited early leaf development triggered by the warm spring. However, yellow birch and American beech have more conservative growth strategies and were largely unaffected by the late spring frost. In contrast, sugar maples responded strongly to warmer temperatures and experi- enced widespread frost damage that resulted in leaf loss and delayed canopy development. Late spring frost events may therefore provide a competitive advantage for yellow birch and American beech at the expense of sugar maple. Results from satellite remote sensing confirm that frost damage was widespread throughout the region at higher elevations (>500 m). The frost event is estimated to have reduced gross ecosystem productivity by 70–153 g C m␣2, or 7–14% of the annual gross productivity (1061 ± 82 g C m␣2) across 8753 km2 of high-elevation forest. We conclude that frost events following leaf out, which are expected to become more common with climate change, may influence both forest composition and ecosystem productivity.

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... These extreme events often have larger and more adverse impacts on ecosystems and society than changes in mean climate alone (Ciais et al., 2005;Reichstein et al., 2013;Orsenigo et al., 2014;Frank et al., 2015). For example, heat waves, droughts, and spring frost defoliations have resulted in large declines in forest carbon uptake in Europe and North America (Ciais et al., 2005;Hufkens et al., 2012). Furthermore, ecosystem responses to extreme climate events often extend well beyond the duration of the event (Frank et al., 2015). ...
... However, earlier leaf out may also increase susceptibility to a late spring frost that causes defoliation of affected trees. While these events typically trigger production of a second cohort of leaves, the effective start of the growing season is delayed, which has the potential to reduce tree growth and forest canopy carbon uptake in temperate broadleaf forests (Hufkens et al., 2012;Wiley et al., 2017;Rubio-Cuadrado et al., 2021). ...
... Because mean leaf-level rates of oak tree photosynthesis were statistically the same in 2020 and 2021, even small changes in LAI from the defoliation event could have impacted total canopy carbon uptake. Delayed leaf expansion and reductions in maximum LAI following a defoliating spring frost have been estimated to reduce gross primary production in a northern hardwood forest by 7-14% (Hufkens et al., 2012). To the extent that defoliation in the oak dominated forests we studied impacted the timing of full leaf expansion and reduced LAI, we might expect the response of total canopy carbon uptake to be similar to what Hufkens et al. (2012) estimated for a northern hardwood forest. ...
Article
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Spring frosts can defoliate trees, reduce canopy carbon assimilation, and alter interspecific competition dynamics. These events may become more common with climate change, but our understanding of the associated ecological impacts is limited by the stochastic nature of their occurrences. In 2020, a late spring frost defoliated oak (Quercus spp.), but not co-occurring maples (Acer spp.) across temperate broadleaf forests of the Hudson Highlands in southern New York State, U.S.A. Defoliation impacted 60% of this region’s forests and delayed full leaf expansion of oaks by ∼17 days. We used this event as an opportunity to advance understanding of how leaf-level physiology, radial growth, and interspecific competition dynamics of mature trees respond to frost-induced defoliation. We quantified leaf-level photosynthetic capacity, stomatal conductance, and water-use efficiency (WUE), as well as basal area increment of defoliated red oak (Q. rubra) trees and non-defoliated red maple (A. rubrum) trees in 2020 (“defoliation year”) and 2021 (“reference year”). Oak defoliation provided red maple trees with a competitive edge in terms of photosynthetic capacity early in the growing season. However, the second cohort of red oak leaves that developed following defoliation had photosynthetic capacities that were 3–4 times higher than red maple trees by the second half of the growing season, likely facilitated by higher rates of stomatal conductance. The growing season mean photosynthetic capacities for the defoliation year were significantly higher for red oaks than red maples. Red oak basal area increment tended to be higher than red maple during both the defoliation and reference years. For both species basal area increment was significantly higher during the reference year than defoliation year, but the reasons remain unclear. Taken together, these findings demonstrate that temporal patterns of photosynthesis in temperate broadleaf forests are altered by defoliation events, but enhanced photosynthetic capacities of second cohort leaves can reduce the negative effects of delayed leaf expansion and mitigate competitive advantages conferred to undefoliated co-occurring tree species. We suggest that understanding a tree species’ ability to compensate for frost-induced defoliation is essential to accurately predict effects of extreme climate events on tree competition dynamics and ecosystem processes.
... An earlier phenology characterized as advanced sprouting and flowering increases the risk of frost damage on newly growing organs, which included sprouted buds, dehardened twigs, unfolded leaves, blooming flowers, and young fruits (Rodrigo 2000, Park 2016, Chmielewski et al. 2018, Vitasse et al. 2018. The loss of carbon (C) in a false spring was unlikely to be offset by post-spring continuous growth under a warming climate (Awaya et al. 2009, Hufkens et al. 2012, Chamberlain et al. 2020. The yearly productivity, however, may have been impaired in forests since the establishment of a late spring frost (Bascietto et al. 2018). ...
... To determine characteristics of a false spring depends on two conditions of dynamic change of temperature. The first condition is achieved with a significant thermal time that advances budburst and flower onset (Hufkens et al. 2012). A certain amount of days with thermal ranges between 0 and 5°C is required for bud burst and leaf unfolding closely following chilling (Cannell & Smith 1986, Heide 1993. ...
... A chronicle of ten years is long enough for analyzing and identifying temperature characters for a late spring frost event (Principe et al. 2017). Current literature indicates a gross period of late spring frost usually happens from a start on 1 March (61 days of year) to 30 April (121 days of years), when it is a time with frequent temperature fluctuations of alternative warming and freeze (Gu et al. 2008, Augspurger 2009, Hufkens et al. 2012, Weather of China 2020. Daily highest and lowest temperature points were recorded to describe the dynamics of extreme temperature in historical spring times as it was employed in former studies (Hufkens et al. 2012, Vitasse et al. 2018. ...
Article
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Information is highly scarce about the possible effect of a late spring frost on physiological response of buds in ornamental trees. In this study, spring temperature of Changchun at Northeast China was recorded to identify the characteristics of a false spring by detecting extraordinary warming and sudden freeze in early April of 2017. Buds of six local ornamental tree species were investigated for their dynamics in biomass, non-structural carbohydrates, frost resistance on days of 7, 14, 21, and 28 April 2017. According to a comparison with spring temperature records historically from 2007 to 2016, a false spring was determined. Black pine (Pinus tabuliformis var. mukdensis) had greater bud biomass than apricot (Prunus sibirica L.). Peach (Prunus persica L. var. persica f. rubro-plena Schneid.) reserved greater non-structural carbohydrate content in post-chilling buds than black pine, and apricot and willow (Salix babylonica L.) had greater soluble sugars and starch contents in buds, respectively. Cumulative number of days with temperature below 12°C had a negative relationship with relative conductance in sorbus (Sorbus pohuashanensis [Hance] Hedl.). Chokecherry (Padus virginiana ‘Canada Red’) had greatest bud starch content on 21 April. Overall, a late spring frost imposed interruption on carbohydrate metabolism rather than direct damage on buds of ornamental trees before late April. Advanced warming induced more pronounced negative impact of a false spring than the sudden decline of minimum temperature.
... Physiological and geological effects of climate change on sugar maple trees were compiled from an evaluation of The New England Regional Assessment of the Potential Consequences of Climate Variability and Change (Lauten et al. 2001), a study of sugar maple's geographical variation of reproductive capacity (Graignic et al. 2014), and reviews of the biophysical responses of plants to climate change effects (Blackwell 2020;Hufkens et al. 2012). ...
... Warmer minimum temperatures in January allow for thawing of snowpack, leading to a reduced snowpack volume and a higher occurrence of soil freezing, causing damage to roots (Oswald et al. 2018). Warmer winter temperatures also present a more frequent spring frost, triggering budbreak 2-3 weeks earlier (Hufkens et al. 2012). The more frequent frost causes leaf-dieback of the initial leaves, forcing sugar maples to expend extra energy to regrow leaves in June and reducing the tree's growing season (Hufkens et al. 2012). ...
... Warmer winter temperatures also present a more frequent spring frost, triggering budbreak 2-3 weeks earlier (Hufkens et al. 2012). The more frequent frost causes leaf-dieback of the initial leaves, forcing sugar maples to expend extra energy to regrow leaves in June and reducing the tree's growing season (Hufkens et al. 2012). ...
Thesis
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Intensifications in seasonal temperature variation, attributed to increased greenhouse gas concentrations, are having an adverse effect on the forest composition of the northeastern United States. New York’s temperate broadleaf forests are undergoing alterations including a decline of the native sugar maple (Acer saccharum), as the species’ population begins to decrease or migrate north to remain within its hospitable temperature zone. By the end of the 21st century, sugar maple’s presence in New York forests will be highly altered from its historical range and population size. The reduction of this species will incur negative effects on New York’s forest ecology, maple-based cultural activities and organizations, and sectors of the local economy. This study examines the primary causes of sugar maple decline and the subsequent ecological, social, and economic impacts of climate change on sugar maple. Potential management preparations, adaptations, and resources to minimize disruptions from sugar maple decline will also be included in this study.
... Phenology is a key component of fitness. Thus, a mismatch between plant phenology and local climate can affect the productivity of crops and forests and carbon uptake of terrestrial ecosystems (Augspurger, 2009;Hufkens et al., 2012), and challenging our strategies of sustainable resource management (Keenan, 2015;Marquis et al., 2022). ...
... Plants from cold and boreal ecosystems are advancing spring phenology (growth reactivation and flowering) due to climate warming (Menzel et al., 2006). The lengthening of the growing season could increase carbon uptake and forest productivity (Keenan et al., 2014), but an early growth reactivation can also expose the young tissues to late frost events (Hufkens et al., 2012). The impact of frosts on plants involves several factors, including a change in frost hardiness during bud break (Bigras and Hébert, 1996). ...
... As late frost damage can have dramatic consequences on plant growth and survival (Allevato et al., 2019;Girardin et al., 2022), late frost occurrence is expected to affect timing of bud break that is critical to determine local adaptation (Kreyling et al., 2014) and species distribution (Kollas et al., 2014). Populations with different sensitivity to climate have different bud phenology and risk of being exposed to late frosts and, consequently, different survival strategies to adapt to the local climate (Hufkens et al., 2012;Marquis et al., 2020a). In the long term, the reduction in fitness caused by recurrent frost damage could change the composition of forest ecosystems by lowering competitiveness and increasing mortality of phenotypes with early bud break and favoring the establishment of those with late bud break (Hufkens et al., 2012). ...
Article
Global warming advances bud break, mismatches plant phenology from the local climate, and exposes the developing leaves to higher risks of frost damage. Bud break of sugar maple [Acer saccharum (Marsh.)], a species included in recent programs of assisted migration, is sensitive to nighttime spring temperatures. This suggests a link between frost events and leaf development. In this study, we raise the hypothesis that late frost is an evolutionary driver of growth reactivation in sugar maple provenances. We investigated the ecotypic variation of bud phenology in 30 provenances planted in two common gardens within and at the northern limit of the species range, in the Province of Quebec, Canada. Eight phases of bud break were assessed twice a week during 2020 on 252 and 272 seedlings in southern and northern sites, respectively. In the southern site, bud break occurred in May, starting on average 12 days earlier and ending 3 days earlier compared to the northern site. Logistic regression was used to estimate the probability of late frost and the results showed that regions located in the north, at higher elevations, and along the northeastern coast of the native maple range showed the latest occurrences of frost events in spring. This pattern mirrored the timing of bud break. When planted in the same common garden, provenances originating from sites with later spring frosts leafed out earlier. Such differences were maintained across the eight bud phenological phases and between the two common gardens, which indicates a similar response of the provenances to changing growing conditions. To avoid frost damage to sugar maple plantations, assisted migration should account for phenotypic traits in bud phenology, ensuring that the frost regime at the origin of the provenances is compatible with that of plantations.
... Thus, an earlier development of the vegetation in spring induced by warming may put plants at a higher risk of being damaged by frosts, particularly in Europe and Asia where tree species have lower freezing resistances (compared to North America) and are generally more responsive to temperature (Liu et al., 2018;Walde et al., 2022;Zohner et al., 2020). In fact, more damaging spring frosts have been observed in recent years, precisely because of the increased amount of heat energy that preceded leafout (Augspurger, 2009;Hufkens et al., 2012;Sangüesa-Barreda et al., 2021;. For instance in April 2017, after an extraordinary warm period in Western and Central Europe, the sudden arrival of cold air masses from the Arctic and the subsequent freezing events caused dramatic losses in fruit production in Central Europe and the amount of accumulated heat at the time of the frost was unprecedented in some stations since the beginning of meteorological records . ...
... Large-scale defoliations due to frost can have a significant impact on forest ecosystem functioning by affecting carbon uptake (Hufkens et al., 2012), water relations (Bréda & Granier, 1996) and by altering nutrient cycles (Estiarte & Peñuelas, 2015). In addition, populations of a vast variety of animal and fungal species, which depend on leaf tissue, fruits and seeds, may decline when frost damages occur (i.e. ...
Article
1. Phenological shifts in response to changing climatic conditions is a key acclimation process for the persistence of perennial plants in temperate and boreal climates. The optimal time to leaf‐out is the result of an evolutionary processes determined by the trade‐off between minimizing the risk of freezing damages and herbivory pressure while maximizing resource uptake to increase competitiveness against the other plants. 2. We quantified the penalty exerted by frost exposure at the time of leaf emergence on plant development (reduction in leaf area, canopy duration, and growth) over the potential gains without frost (increased biomass and non‐structural carbohydrate reserves), depending on when leaf‐out occurs. To this purpose, we exposed 960 saplings of four temperate deciduous tree species with contrasting cold hardiness to two frost intensities shortly after leaf emergence, which was artificially induced at four occasions to reflect the whole range of natural leaf‐out dates. 3. One year above‐ground biomass (AGB) increments following the frost revealed a clear ranking among the species depending on their strategy to cope with damaging frosts. Prunus avium (‐41% of AGB‐increment compared to control saplings) resprouted from the stem base, Quercus robur (‐62%) rapidly produced new leaves from dormant reserve buds, Fagus sylvatica (‐98%) showed highest chlorophyll content in autumn and delayed senescence together with Carpinus betulus (‐105%), which overcompensated NSC reserves after the growing season but showed highest mortality (up to 32%). In all species, NSC reserves recovered rapidly their initial stage at the expense of growth. 4. The timing of leaf‐out (advanced and delayed artificially) significantly affected the performance and recovery (regreening and growth) of both frozen and non‐frozen saplings, with the lowest performance found at the most delayed leaf‐out date. We propose that the potential to recover from frost damages is an important component of a tree's performance, particularly at the juvenile stage. The ability to recover may become even more decisive in the future with the predicted increase of false springs in many extra‐tropical regions.
... For example, the economic loss for agricultural crops after the 2007 late-spring frost in the U.S. was up to $112 million, with fruit crop loss of $86 million (Warmund et al., 2008). Further examples of late-spring frost damage include 1995 in France (Ningre & Colin, 2007); 2010 in the NE of U.S.A. (Hufkens et al., 2012); and 2011 and 2016 in Switzerland, south Germany, and northeastern France (Kreyling et al., 2012;. ...
... Late-spring frost can also exert substantial impacts on trees, including increased mortality, altered morphology, decreased growth, changes in primary nutrients for insects (e.g., N), and changes in leaf chemical defences (St. Clair et al., 2009;Hufkens et al., 2012;Man et al., 2013). Consequently, late-spring frost may also influence the susceptibility of plants to insects or pathogens, although the number of studies that have examined the connection between freeze damage and insect susceptibility in trees is limited. ...
Article
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• Global climate change affects the frequency of extreme weather events that can influence plant–insect interactions. • We evaluated how the late‐spring frost and severe drought that occurred in Spain in 2017 affected interactions between the invasive gall insect, Dryocosmus kuriphilus, and the native tree, Castanea sativa. We assessed effects on insect survival, fertility, population growth, and effects through changes in tree palatability and in other pests and pathogens. • Late‐spring frost reduced D. kuriphilus to 25–40% of previous abundance. Wasp populations recovered rapidly (>7‐fold in 3 years), consistent with density‐dependence in population dynamics. • Larvae affected by freeze or drought were smaller. Female fecundity was affected by the freeze 1 year later. • Late‐spring frosts and severe drought affected leaf size and physiology. Water content was higher within galls, but nitrogen was higher within galls in non‐freeze plots after weather conditions improved. • Freezing also influenced the secondary chemistry of leaves. Phenol concentrations were lower, and terpenes higher, in frozen plots, while condensed tannins remained the same. Condensed tannins were reduced to half in the drought year. • Freezing had limited effects on damage from other pests and pathogens. • Our work expands understanding of how climate and weather affects forest pests.
... The results of our study showed immediate, negative effects on vegetative phenophases-although these effects were not significant. Although our findings were consistent with numerous other studies, e.g., [77][78][79][80][81][82][83][84][85], other studies found contradictory results, e.g., [86][87][88]. The reason for these differences may have been related to the dependence of the phenological cycle on environmental factors and stressors, such as drought, fire, soil erosion, and soil properties, as well as climatic and anthropogenic (e.g., land use and land cover change). ...
Article
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Wildfire has significant impact on plant phenology. The plants’ phenological variables, derived from time series satellite data, can be monitored and the changes in satellite imagery may be used to identify the beginning, peak, and end of the growing season. This study investigated the use of remote sensing data and land surface phenology (LSP) parameters to evaluate the impacts of fire. The LSP parameters included the start of growing season (SOS), the length of the growing season (LOS), the end of the growing season (EOS), maximum greenness of the season (Gmax), and minimum greenery in the season (Gmin) in the fire-impacted, semiarid oak forests of Iran. These LSP parameters were extracted from multitemporal normalized difference vegetation index (NDVI) and enhanced vegetation index (EVI2) data, acquired from MODIS sensor images in Zagros of the Ilam province in western Iran. By extracting LSP indices from the NDVI and EVI2 data, the indices were compared between burned forest areas, areas surrounding the burned forests, and unburned areas and for timesteps representing pre-fire, fire (i.e., year of fire), and post-fire (i.e., 2 years) conditions. It was found that for the burned area, there were significant differences in Gmax and the day that Gmax occurred. Furthermore, there was also a significant difference in Gmin between the pre- and post-fire conditions when NDVI was used and a significant difference between Gmax when EVI2 was used. The results also showed that in both time series there was a significant difference between the burned and control area in terms of Gmax. In general, the results showed that the fire had a negative effect on LSP, but in the two years after the fire, there were signs of forest restoration. This study provides necessary information to inform forest and resource conservation and restoration programs.
... Different species have separate growth strategies, tolerances, and resilience, resulting in various levels of impact. For example, conservative growth strategies render yellow birch and American beech largely unaffected by late spring frosts, while sugar maple suffers from frost damage, resulting in leaf loss and delayed crown development [64]. In addition, European beech showed low resistance to late frost events but high resilience in radial growth [65]. ...
Article
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The response of vegetation spring phenology to climate warming has received extensive attention. However, there are few studies on the response of vegetation spring phenology to extreme climate events. In this study, we determined the start of the growing season (SOS) for three vegetation types in temperate China from 1982 to 2015 using the Global Inventory Modeling and Mapping Study’s third-generation normalized difference vegetation index and estimated 25 extreme climate events. We analyzed the temporal trends of the SOS and extreme climate events and quantified the relationships between the SOS and extreme climate events using all-subsets regression methods. We found that the SOS was significantly advanced, with an average rate of 0.97 days per decade in China over the study period. Interestingly, we found that the SOS was mainly associated with temperature extremes rather than extreme precipitation events. The SOS was mainly influenced by the frost days (FD, r = 0.83) and mean daily minimum temperature (TMINMEAN, r = 0.34) for all three vegetation types. However, the dominant influencing factors were vegetation-type-specific. For mixed forests, the SOS was most influenced by TMINMEAN (r = 0.32), while for grasslands and barren or sparsely vegetated land, the SOS was most influenced by FD (r > 0.8). Our results show that spring phenology was substantially affected by extreme climate events but mainly by extreme temperature events rather than precipitation events, and that low temperature extremes likely drive spring phenology.
... Early studies suggested that rapidly differentiating and actively growing leaves and flowers are most susceptible to frost (Augspurger, 2009;Hufkens et al., 2012;Sakai and Larcher, 1987). So the declining temperature sensitivity of spring phenology was beneficial for the plants since it reduced the risk of late spring frost damage (Fu et al., 2015). ...
Article
Changes in the rate of spring green-up (RSP) caused by climate change can alter biosphere-atmosphere exchange of carbon, energy, and water cycles. Although the response of RSP to environmental conditions has been frequently discussed, little is known about the dynamic relationship between RSP and environmental variables over time. We investigated the interannual relationships between RSP and various environmental variables using the Ensemble Empirical Mode Decomposition (EEMD) detrending approach and partial correlation analyses. We then used 15-year moving windows to analyze the effects of accumulated green-up period temperature (AGT) on RSP and its temporal variation from 1982 to 2016.We found that: (1) AGT had a positive influence on RSP in 81.08% of the entire study regions and AGT dominated the interannual variation of RSP in 31.31% all pixels; (2) The percentage of moisture-controlled (vapor pressure deficit (VPD) and soil moisture) pixels was 28.71%, indicating that preseason moisture, especially vapor pressure deficit (negative correlations in 65.50% pixels), played critical roles in limiting RSP; (3) The linkage between RSP and AGT (RRSP–AGT) was significantly weakening over the Northern Extratropics with intensity decrease of −0.0017 year⁻¹ (p < 0.05) and extent reduction of −0.4 × 10⁵ km² year⁻¹ (p < 0.05), suggesting a decreasing influence of AGT on foliar development. (4) The decline in RRSP–AGT coincided with decreased chilling exposure (CE) and enhanced VPD during the preseason over the Northern Extrotrapics and Eurasia, but it only may be related to changes in VPD in the North America. Our findings highlighted the importance of the dynamic response of vegetation spring green-up to ongoing warming, for a better understanding of the vegetation seasonal cycle under climate change.
... In addition, climate warming could redistribute the optimal species growing in different regions, causing northward expansion of forest tree line and promote ecosystem species competition and succession ( Huang et al., 2017 ), and then influence the regional landcover type, land surface characteristics and soil carbon component ( Peñuelas et al., 2009 ). These phenology-related changes could alter the interactions between trophic levels and results in niche mismatch, biodiversity shifts, which could change the original carbon transport path of the ecosystem, even truncate some of the path, and consequentially affect the terrestrial carbon balance ( Augspurger, 2009 ;Hufkens et al., 2012 ;Trumbore et al., 2015 ). ...
Article
The Earth is experiencing unprecedented climate change. Vegetation phenology has already showed strong response to the global warming, which alters mass and energy fluxes on terrestrial ecosystems. With technology and method developments in remote sensing, computer science and citizen science, many recent phenology-related studies have been focused on macrophenology. In this perspective, we 1) reviewed the responses of vegetation phenology to climate change and its impacts on carbon cycling, and reported that the effect of shifted phenology on the terrestrial carbon fluxes is substantially different between spring and autumn; 2) elaborated how vegetation phenology affects ecohydrological processes at different scales, and further listed the key issues for each scale, i.e. focusing on seasonal effect, local feedbacks and regional vapor transport for individual, watershed and global respectively); 3) envisioned the potentials to improve current hydrological models by coupling vegetation phenology-related processes, in combining with machine learning, deep learning and scale transformation methods. We propose that comprehensive understanding the climate-macrophenology-hydrology interactions are essential and urgently needed for enhancing our understanding of the ecosystem response and its role in hydrological cycle under future climate change.
... Although LSP observations can be influenced by several factors (e.g. spatial resolution, understorey, time series processing, metrics calculation; de Beurs and Henebry 2005), repeated evidence showed their ability to catch forest canopy dynamics (Liang et al. 2011;Hufkens et al. 2012;Richardson et al. 2018). ...
Article
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Calibrating land surface phenology (LSP) with tree rings is important to model spatio-temporal variations in forest productivity. We used MODIS (resolution: 250 m) NDVI, WDRVI and EVI series 2000-2014 to derive LSP metrics quantifying phenophase timing and canopy photosynthetic rates of 26 European beech forests covering a large thermal gradient (5-16 °C) in Italy. Average phenophase timing changed greatly with site temperature (e.g. growing season 70 days longer at low- than high-elevation); average VI values were affected by precipitation. An annual temperature about 12 °C (c. 1100 m asl) represented a bioclimatic threshold dividing warm from cold beech forests, distinguished by different phenology-BAI (basal area increment) relationships and LSP trends. Cold forests showed decreasing VI values (browning) and delayed phenophases and had negative BAI slopes. Warmer forests tended to increase VI (greening), and positive BAI slopes. NDVI peak, commonly used in global trend assessments, changed with elevation in agreement with changes in wood production. A cross-validation modelling approach demonstrated the ability of LSP to predict average BAI and its interannual variability. Merging sites into bioclimatic groups improved models by amplifying the signal in growth or LSP. NDVI had highest performances when informing on BAI trends; WDRVI and EVI were mostly selected for modelling mean and interannual BAI. WDRVI association with tree rings, tested in this study for the first time, showed that this VI is highly promising for studying forest dynamics. MODIS LSP can quantify forest functioning changes across landscapes and model interannual spatial variations and trends in productivity dynamics under climate change.
... B.S.P.] is a relevant model species to study this problem. The current scenario of increasing risk of late frost would entail negative impacts on growth, carbon uptake and survival of boreal species by loss of photosynthetic and reproductive tissue (Hufkens et al., 2012) and potentially increase the susceptibility to other stressors, such as drought . ...
Article
Under climate change, the increasing occurrence of late frost combined with advancing spring phenology can increase the risk of frost damage in trees. In this study, we tested the link between intra-specific variability in bud phenology and frost exposure and damages. We analysed the effects of the 2021 late frost event in a black spruce (Picea mariana (Mill.) BSP) common garden in Québec, Canada. We hypothesized that the timing of budbreak drives the exposure of vulnerable tissues and explains differences in frost damage. Budbreak was monitored from 2015 to 2021 in 371 trees from five provenances originating between 48° and 53° N and planted in a common garden at 48° N. Frost damages were assessed on the same trees through the proportion of damaged buds per tree and related to the phenological phases by ordinal regressions. After an unusually warm spring, minimum temperatures fell to -1.9°C on May 28 and 29, 2021. At this moment, trees from the northern provenances were more advanced in their phenology and showed more frost damage. Provenances with earlier budbreak had a higher probability of damage occurrence according to ordinal regression. Our study highlights the importance of intra-specific variability of phenological traits on the risk of frost exposure. We provide evidence that the timings of bud phenology affect sensitivity to frost, leading to damages at temperatures of -1.9°C. Under the same conditions, the earlier growth reactivation observed in the northern provenances increases the risks of late frost damage on the developing buds. This article is protected by copyright. All rights reserved.
... Additionally, nighttime warming reduces the frost risk in spring ( Figure S11). Newly developed leaves lack the structural integrity to prevent frost damage (Hufkens et al., 2012), and frost during the early stage of vegetation greenup will directly affect subsequent leaf development and vegetation growth throughout the year (Deng et al., 2020). ...
Article
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Recent studies have revealed that the influence of diurnal temperature on spring phenology is asymmetric, and the faster nighttime warming in the Northern Hemisphere (NH) has a complex impact on spring phenology. Our understanding from the sensitivity of the start of the growing season (SOS) to daytime (ST_daytime) and nighttime temperatures (ST_nighttime) has urgently needs to be improved. In this study, the SOS sensitivity to diurnal temperature in the middle and high latitudes of the NH (>30°N) from 1982‐2015 is estimated. The results indicate that although SOS showed stronger sensitivity to daytime than nighttime temperature in most parts of the study areas, the influence of daytime temperature on SOS is decreasing, while the influence of nighttime temperature on SOS is increasing. The variations in ST_daytime and ST_nighttime along the latitude gradient were significantly correlated with the warming rate of the preseason diurnal temperature (p<0.01). The SOS between 40°N and 70°N was more sensitive to daytime temperature, while ST_nighttime was higher than ST_daytime at other latitudes due to topography and rapid nighttime warming. On the altitude gradient, the SOS was more sensitive to daytime temperature in areas below 800 m and 2000‐4000 m. ST_nighttime exceeded ST_daytime at other altitudes owing to nighttime warming relief of the severe restrictions on phenological processes and the reduction in frost risk. To reach a comprehensive characterization of the interaction between vegetation and climate systems, the current study suggests more investigation on the response of SOS to diurnal temperature on large scales. This article is protected by copyright. All rights reserved.
... Coping with frost requires adapted behaviours or elaborate physiological adaptations for both ecto-and endothermal organisms, and especially for non-migrating life forms that cannot escape, such as plants. Frost change frequency carries information about the occurrence frequency of freezing and thawing events and -indirectly -about their duration, both of which are crucial constraints determining the best-suited adaptation strategies, see e.g., Hufkens et al. (2012). We used a Bspline interpolation S(tasmax, t) and S(tasmin, t) to get both daily minimum (tasmini) and maximum (tasmaxi) near-surface 2 255 m air temperatures from monthly values, with t the sequence of Julian days marking the middle of each month, i.e., [349,15,45,74,105,135,166,196,227,258,288,319,349,15]. ...
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A multitude of physical and biological processes on which ecosystems and human societies depend are governed by climatic conditions. Understanding how these processes are altered by climate change is central to mitigation efforts. Based on mechanistically downscaled climate data, we developed a set of climate-related variables at yet unprecedented spatiotemporal detail as a basis for environmental and ecological analyses. We created gridded data for near-surface relative humidity (hurs), cloud area fraction (clt), near-surface wind speed (sfcWind), vapour pressure deficit (vpd), surface 15 downwelling shortwave radiation (rsds), potential evapotranspiration (pet), climate moisture index (cmi), and site water balance (swb), at a monthly temporal and 30 arcsec spatial resolution globally, from 1980 until 2018 (time-series variables). At the same spatial resolution, we further estimated climatological normals of frost change frequency (fcf), snow cover days (scd), potential net primary productivity (npp), growing degree days (gdd), and growing season characteristics for the periods three shared socioeconomic pathways (SSP126, SSP370, 20 SSP585) and five Earth system models (projected variables). Time-series variables showed high accuracy when validated against observations from meteorological stations. Projected variables were also highly correlated to observations, although some variables showed notable biases, e.g., snow cover days (scd). Together, the CHELSA-BIOCLIM+ data set presented here (https://doi.org/10.16904/envidat.332, Brun et al., 2022) allows improving our understanding of patterns and processes that are governed by climate, including the impact of recent and future climate changes on the world's ecosystems and 25 associated services to societies.
... Atmospheric phenomena, especially frost, are among the costliest natural hazards in the world (Guan et al., 2015), which have a massive effect on agriculture (Xiao et al. 2018), horticulture (Moonen et al. 2002;Kamali et al. 2008;Chmielewski et al. 2018), and forestry (Gu et al. 2008;Hufkens et al. 2012;Sangüesa-Barreda et al. 2021;Lhotka Brönnimann, 2021;Kollas et al. 2014;Meier et al. 2018). Frost is defined in various ways, including a decline in temperature below the freezing point of water during the growing period, a reduction in temperature below 0 °C during the positive numbers of temperature in 24 h, and a fall in air temperature below 0° C during a 24-h period with a maximum positive temperature (Niedzwied, 2003). ...
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Occurrence of extreme climatic phenomena such as frost will cause significant risks and costs to many sectors, especially agriculture, horticulture, and forests. Frost will cause the worst damage when it occurs at the critical stages of crops, especially in spring. The frost phenomena are one of the important climatic and environmental hazards that cause a lot of damage to the agricultural sector of Iran every year. In this respect, the present study intends to highlight the projection of late spring rost by global circulation models (GCMs) from Coupled Model Intercomparison Project Phase 6 (CMIP6). For this purpose, minimum temperature data of 17 synoptic stations were used in the period 1985-2014 in cold regions of Iran. For projecting the changes of LSF, the ACCESS-ESM1-5 and Nor-ESM2-LM Models were used under three (Shared Socioeconomic Pathway (SSP)) scenarios SSP1-2.6, SSP2-4.5, and SSP5-8.5 for the next three periods (i.e., 2020-2049, 2050-2079, and 2080-2099). Then, the changes were compared to the historical period (1985-2014). The root mean square error (RMSE), mean bias error (MBE), correlation coefficient (CC), and Nash-Sutcliff efficiency (NSE) indices evaluated the models' performances. The results revealed that the latest and earliest dates of LSF during the base period occurred in the western and central parts of Iran, respectively. The model evaluation indicated that the performance of ACCESS-ESM1-5 (MBE = 0.3, CC = 0.87, and NSE = 0.68) exhibited a higher accuracy than the NorESM2-LM model. Based on both GCM under all three SSP scenarios, the projection of changes in future periods (compared to the base period) indicated that the date of occurrence of LSF would be earlier than the base period, with the highest and lowest changes projected based on NorESM2-SSP5-8.5 and ACCESS-ESM1-5-SSP1-2.6 in Arak, Isfahan, Khorramabad, Sabzevar, Shahrekord, and Shahroud stations. In general, depending on the model and climate scenario, the LSF phenomenon occurs earlier or later in cold regions of Iran, and its changes would be between − 76 and + 19 days in the future period.
... Meanwhile, air Gordo and Sanz (2009) suggested the 1 • C warming in early spring was observed to bring an advance of spring phenology by up to 7 days for the Spanish populations of European beech. Hufkens et al. (2012) indicated that the low temperature events in the late spring of 2010 contributed to a 14% ∼ 33% decreasing in the leaf area index over the northeastern United States. Therefore, it is reasonable to examine whether the winter North Atlantic SST influences the spring NDVI by regulating the local temperature. ...
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In this study, the single value decomposition method was used to analyze the covariation between the winter (December, January, and February) North Atlantic sea surface temperature (SST) and spring (April and May) normalized difference vegetation index (NDVI) over mid‐high‐latitude Eurasia for the period 1982–2015. The results show that the first paired mode explains 32.8% of the total squared covariance, and the correlation of time coefficients is +0.67 (p < 0.01), suggesting that the meridional tripole structure of the winter North Atlantic SST is significantly associated with the three zonally anomalous centers of the spring mid‐high‐latitude Eurasian NDVI. Surface air temperature (T2m) is the most important factor related to spring NDVI changes. The tripole pattern of the winter North Atlantic SST can persist until the following spring, triggering an eastward anomalous Rossby wave train in the middle troposphere, which features two negative geopotential height anomalies centers near northwestern Europe and eastern Siberia and one positive center over central Eurasia. Surface temperature anomalies are well consistent with these anomalous pressure centers. This implies that the winter North Atlantic SST could be a robust driver and potential precursor for spring vegetation prediction over the mid‐high latitudes of Eurasia.
... In a hardwood Mediterranean forest in southwest Australia, tree canopy frost damage increased with declining elevation because of cold-air pooling (Matusick et al., 2014). In temperate deciduous forests, frost damage could be especially devastating for species like sugar maple, which tends to leaf out early in the season to optimize carbon gain but is particularly vulnerable to frost damage and temperature anomalies (Hufkens et al., 2012), as well as for saplings that experience earlier budbreak than conspecific canopy trees (Augspurger, 2009;Pederson et al., 2014). In other cases, deciduous species could respond to cold stress in these areas by delaying spring phenology (e.g., later budburst and leaf emergence) and shedding leaves earlier in fall (Vitasse et al., 2014), shortening the growing season and window for carbon capture. ...
Article
Cold‐air pooling is a global phenomenon that frequently sustains low temperatures in sheltered, low‐lying depressions and valleys and drives other key environmental conditions, such as soil temperature, soil moisture, vapor pressure deficit, frost frequency, and winter dynamics. Local climate patterns in areas prone to cold‐air pooling are partly decoupled from regional climates and thus may be buffered from macroscale climate change. There is compelling evidence from studies across the globe that cold‐air pooling impacts plant communities and species distributions, making these decoupled microclimate areas potentially important microrefugia for species under climate warming. Despite interest in the potential for cold‐air pools to enable species persistence under warming, studies investigating the effects of cold‐air pooling on ecosystem processes are scarce. Because local temperatures and vegetation composition are critical drivers of ecosystem processes like carbon cycling and storage, cold‐air pooling may also act to preserve ecosystem functions. We review research exploring the ecological impacts of cold‐air pooling with a focus on vegetation, and then present a new conceptual framework in which cold‐air pooling creates feedbacks between species and ecosystem properties that generate unique hotspots for carbon accrual in some systems relative to areas more vulnerable to regional climate change impacts. Finally, we describe key steps to motivate future research investigating the potential for cold‐air pools to serve as microrefugia for ecosystem functions under climate change.
... adjusting phenology to new climatic conditions) can lead to greater growth and/or reproduction in some species (Cleland et al., 2012). However, early leaf out can also be problematic if it occurs during a period of warming followed by a hard frost, known as a false spring (Inouye, 2008;Hufkens et al., 2012). In the end, what matters is the 'match' between phenology and the environment, and this 'match' is influenced by species-specific differences in anatomy and physiology (Cooke et al., 2012;Panchen et al., 2014;Flynn & Wolkovich, 2018). ...
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There is a long‐standing idea that the timing of leaf production in seasonally cold climates is linked to xylem anatomy, specifically vessel diameter because of the hydraulic requirements of expanding leaves. We tested for a relationship between the timing of leaf out and vessel diameter in 220 plants in three common gardens accounting for species’ phylogenetic relationships. We investigated how vessel diameter related to wood porosity, plant height and leaf length. We also used dye perfusion tests to determine whether plants relied on xylem produced during the previous growing season at the time of leaf out. In all three gardens, there was later leaf out in species with wider vessels. Ring‐porous species had the widest vessels, exhibited latest leaf out and relied less on xylem made during the previous growing season than diffuse‐porous species. Wood anatomy and leaf phenology did not exhibit a phylogenetic signal. The timing of leaf out is correlated with wood anatomy across species regardless of species’ geographic origin and phylogenetic relationships. This correlation could be a result of developmental and physiological links between leaves and wood or tied to a larger safety efficiency tradeoff.
... Climate change not only results in elevated mean temperatures in spring, but also increases the risk of late spring frost events, particularly in Europe and Asia [25][26][27]. Late spring frosts are rare but may heavily impact forest ecosystems as they can negatively affect, both directly and indirectly, tree survival, growth, regeneration and stem form [28,29]. For instance, radial growth in central European beech was reduced by 90% due to late spring frosts [30]. ...
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Global change increases the risk of extreme climatic events. The impact of extreme temperature may depend on the tree species and also on the provenance. Ten provenances of Fagus sylvatica L. were grown in a common garden environment in Belgium and subjected to different temperature treatments. Half of the one year old seedlings were submitted to a high thermal stress in the spring of the first year, and all plants were exposed to a late spring frost in the second year. The high-temperature treated plants displayed reduced growth in the first year, which was fully compensated (recovery with exact compensation) in the second year for radial growth and in the third year for height growth. Frost in the spring of the second year damaged part of the saplings and reduced their growth. The frost damaged plants regained the pre-stress growth rate one year later (recovery without compensation). The high temperature treatment in the first year and the frost damage in the second year clearly influenced the phenological responses in the year of the event and in the succeeding year. Little population differentiation was observed among the provenances for growth and for phenological responses. Yet, a southern provenance, a non-autochthonous provenance (original German provenance that was planted in Belgium about a century ago) and a more continental provenance flushed earlier than the local Atlantic provenances in the year of the frost event, resulting in more frost damage. Some caution should therefore be taken when translocating provenances as an anticipation of the predicted climate warming.
... Sugar maple is generally healthy across its range, though declining growth and crown dieback have been noted in selected stands in the last 40 years (Bauce and Allen 1992;Payette et al. 1996). The specific drivers of decline, often termed sugar maple decline disease, are diverse and often locally specific but well studied (Horsley et al. 2002;Hufkens et al. 2012). Given the species' specific environmental (Sullivan et al. 2013;Bauce and Allen 1992;Duchesne et al. 2002) and climate (Gavin 2008) tolerances, climate change is likely to be the largest threat to the species in the future (Bishop et al. 2015). ...
Thesis
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Unabated anthropogenic greenhouse gas emissions have resulted in an unprecedented rise in global atmospheric CO2 (Ca) levels and a perhaps irreversible shift in the dynamics of global carbon cycling. Given that C3 plants are likely to be carbon limited at current Ca levels, there was previous optimism regarding the potential for increased carbon storage and water-use efficiency (WUE) in global forests. To address this question, scientists have turned to tree ring series to investigate long-term trends in growth and resource use in global forests. Unfortunately, our ability to draw accurate conclusions regarding the fate of forests from tree ring studies alone is limited by our inability to control for the multitude of environmental and developmental variables that confound long-term, climate-related signals in tree ring series. In this thesis I use principles of forest ecology to select forest types where long-term changes in growth and WUE can be estimated independently of the effects of stand and individual tree development, namely, chronosequence jack pine and self-replacing sugar maple forests in Ontario. To do so a novel tree ring standardization model is presented that uses tree diameter in the year of ring formation as the primary determinant of the underlying developmental trend. This method is shown to be superior to current models in separating developmental trends from long-term environmental/climatic signals in tree ring series from shade-tolerant species. In chronosequence jack pine stands I show increases in stand-level WUE but progressive growth decline. Water-use efficiency was negatively associated with tree growth, suggesting that warming- and drought-induced stomatal closure has likely led to deviations from expected Ca enhanced growth. In self-replacing sugar maple, I show that the response of neither growth or WUE to increasing Ca is conserved across the site- or landscape-levels. While it is evident that variability in soil moisture controls this response at the site-level, the drivers of variation across the landscape are unclear. Further, high-frequency climate sensitivity is not conserved across stands near the species northern range limit, nor is climate an important predictor of growth in these stands. These findings have important implications for range prediction of the species, as current distribution models are climate driven.
... Although the timing of leaf senescence is controlled to some extent by leaf age (Reich et al. 1992;Lim et al. 2007) and is thus endogenous, environmental factors and biotic interactions can alter the relationship between spring and autumn phenology in several ways. Individuals with early spring phenology may have greater exposure to frost (Bennie et al. 2010;Hufkens et al. 2012) or insect attack (Wesolowski and Rowinski 2008;Jepsen et al. 2011), which are both linked with earlier senescence. Abiotic and biotic factors may also affect phenological correlations by altering photosynthetic or carbon assimilation rates, which are known to influence senescence timing (Zani et al. 2020). ...
Article
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Seasonal life history events are often interdependent, but we know relatively little about how the relationship between different events is influenced by the abiotic and biotic environment. Such knowledge is important for predicting the immediate and evolutionary phenological response of populations to changing conditions. We manipulated germination timing and shade in a multi-factorial experiment to investigate the relationship between spring and autumn phenology in seedlings of the pedunculate oak, Quercus robur, and whether this relationship was mediated by natural colonization of leaves by specialist fungal pathogens (i.e., the oak powdery mildew complex). Each week delay in germination corresponded to about 2 days delay in autumn leaf senescence, and heavily shaded seedlings senesced 5–8 days later than seedlings in light shade or full sun. Within seedlings, leaves on primary-growth shoots senesced later than those on secondary-growth shoots in some treatments. Path analyses demonstrated that germination timing and shade affected autumn phenology both directly and indirectly via pathogen load, though the specific pattern differed among and within seedlings. Pathogen load increased with later germination and greater shade. Greater pathogen load was in turn associated with later senescence for seedlings, but with earlier senescence for individual leaves. Our findings show that relationships between seasonal events can be partly mediated by the biotic environment and suggest that these relationships may differ between the plant and leaf level. The influence of biotic interactions on phenological correlations across scales has implications for understanding phenotypic variation in phenology and for predicting how populations will respond to climatic perturbation.
... SFD is a well-known worldwide risk for overwintering crops, tree species, and some early spring orchards in temperate climate regions [8,50,51] and Mediterranean areas [49,52] at northern latitudes, and also for tea plantations in humid areas around the equator [14]. The distribution characteristics of SFD risk were systematically analyzed at three scales, including temporal scale, spatial scale, and terrain scale, over the MLRYR in China. ...
Article
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Spring frost damage (SFD), defined as the disaster during the period of newly formed tea buds in spring caused by lower temperature and frost damage, is a particular challenge for tea plants (Camellia sinensis), whose capacity to adapt to extreme weather and climate impacts is limited. In this paper, the region of the Middle and Lower Reaches of the Yangtze River (MLRYR) in China was selected as the major tea plantation study area, and the study period was focused on the concentrated occurrence of SFD, i.e., from March to April. By employing the standard lapse rate of air temperature with elevation, a minimum temperature (Tmin) estimation model that had been previously established was used based on reconstructed MYD11A1 nighttime LST values for 3 × 3 pixel windows and digital elevation model data. Combined with satellite-based Tmin estimates and ground-based Tmin observations, the spatiotemporal characteristics of SFD for tea plants were systematically analyzed from 2003 to 2020 in the MLRYR. The SFD risks at three scales (temporal, spatial, and terrain) were then evaluated for tea plants over the MLRYR. The results show that both SFD days at the annual scale and SFD areas at the daily scale exhibited a decreasing trend at a rate of 2.7 days/decade and 2.45 × 104 ha/day, respectively (significant rates at the 0.05 and 0.01 levels, respectively). The period with the highest SFD risk appeared mainly in the first twenty days of March. However, more attention should be given to the mid-to-late April time period due to the occurrence of late SFD from time to time. Spatially, areas with relatively higher SFD days and SFD risks were predominantly concentrated in the higher altitude areas of northwestern parts of MLRYR for both multi-year averages and individual years. Fortunately, in regions with a higher risk of SFD, the distribution of tea plants was relatively scattered and the area was small. These findings will provide helpful guidance for all kinds of people, including government agencies, agricultural insurance agencies, and tea farmers, in order that reasonable and effective strategies to reduce losses caused by spring frost damage to tea plants may be recommended and implemented.
... Previous work in the UK revealed a tendency for host alternating aphids to delay their autumn return journey to primary host plants in warm years (Bell et al., 2015), which might narrow the window of opportunity for mating and egg laying on primary host plants before the onset of winter. Increasing climate variability and extreme weather events might also disproportionately affect host alternating aphids, for example, when a warm spell in the spring induces egg hatch only to be followed by a period of lethally cold low temperatures (Augspurger, 2013;Hufkens et al., 2012). Host alternation might also increase vulnerability to climate change by constraining poleward range expansion as aphids follow their thermal optima, if either winter or summer host plants do not also shift their ranges poleward (Bell et al., 2012). ...
Article
Many animals change feeding habits as they progress through life stages, exploiting resources that vary in space and time. However, complex life histories may bring new risks if rapid environmental change disrupts the timing of these switches. Here, we use abundance times series for a diverse group of herbivorous insects, aphids, to search for trait and environmental characteristics associated with declines. Our meta dataset spanned three world regions and >300 aphid species, tracked at 75 individual sites for 10-50 years. Abundances were generally falling, with median changes of -8.3%, -5.6%, and -0.1% per year in the central USA, northwestern USA, and United Kingdom, respectively. Aphids that obligately alternated between host plants annually and those that were agricultural pests exhibited the steepest declines, relative to species able to persist on the same host plant year-round or those in natural areas. This suggests that host alternation might expose aphids to climate-induced phenology mismatches with one or more of their host plant species, with additional risks from exposure to insecticides and other management efforts. Warming temperatures through time were associated with milder aphid declines or even abundance increases, particularly at higher latitudes. Altogether, while a warming world appeared to benefit some aphid species in some places, most aphid species that had time-sensitive movements among multiple host plants seemed to face greater risk of decline. More generally, this suggests that recent human-induced rapid environmental change is rebalancing the risks and rewards associated with complex life histories.
... Globally, too, time-lapse cameras have been deployed for analysing plant phenophases (Richardson et al. 2018a), crop productivity (Moriondo et al. 2016), natural resource conservation (Bater et al. 2011), experimental ecology (Pennekamp Fig. 2 Classification of pixels into the 'snow' and 'no snow' categories based on their blue digital numbers using the threshold limit (left side) along with photograph of that particular day (right side); amaximum snow cover, bpartial snow cover, and cno snow. and Schtickzelle 2013) and frost damage (Hufkens et al. 2012b). However, the present study is amongst the pioneer works from a global biodiversity hotspot that is highly heterogeneous. ...
Article
The use of time-lapse camera setups for characterizing phenology is fast emerging because of their advantages in offering continuous unbiased data. We therefore installed a camera setup in the Western Himalaya to monitor temporal patterns of Betula utilis phenology and also to document snow cover patterns. Digital images (N = 653) of two growing seasons (2017 and 2018) captured through the setup were used for the same. Images hold information in the red, green and blue channels (RGB) and relative changes in RGB indicate canopy colouration. We categorized the phenophases into greenup, leaf maturity, senescence and dormancy. The RGB analyses revealed that during both the years, greenup in B. utilis started during early May [128th and 124th day of the year (DOY) in 2017 and 2018, respectively] and continued till mid-June when the canopy attained maturity. On the other hand, senescence started in early September and by mid-October, the trees became leafless (288th and 289th DOY in 2017 and 2018, respectively). A four-day earlier greenup and dormancy delayed by one day were noted in 2018 when compared to 2017. Thus, the length of the growing season was five days longer in 2018. The snow cover ratio revealed that snowmelt occurred eigth days earlier in 2018 than in 2017. Though this is preliminary, seasonal phenological patterns are evident and call for continued monitoring of B. utilis. The installed setup will provide continuous long-term data from the Himalayan region which had been lacking until now. The present setup for phenological monitoring is pioneering in the Indian Himalaya and needs to be replicated.
... The risk of false springs was identified early in our understanding of the effects of climate change on species' phenology [13]. Since then, there have been numerous documented instances of freeze-induced plant death and damage due to premature budding and shooting (e.g., in the winters of 2007 [26] and 2010 [36]). ...
Article
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Background Increases in temperature variability associated with climate change have critical implications for the phenology of wildlife across the globe. For example, warmer winter temperatures can induce forward shifts in breeding phenology across taxa (“false springs”), which can put organisms at risk of freezing conditions during reproduction or vulnerable early life stages. As human activities continue to encroach on natural ecosystems, it is also important to consider how breeding phenology interacts with other anthropogenic stressors (e.g., pollutants). Using 14 populations of a widespread amphibian (wood frog; Rana sylvatica ), we compared 1) growth; 2) tolerance to a common wetland contaminant (NaCl); and 3) the ability of tadpoles to acclimate to lethal NaCl exposure following sublethal exposure earlier in life. We evaluated these metrics across two breeding seasons (2018 and 2019) and across populations of tadpoles whose parents differed in breeding phenology (earlier- versus later-breeding cohorts). In both years, the earlier-breeding cohorts completed breeding activity prior to a winter storm and later-breeding cohorts completed breeding activities after a winter storm. The freezing conditions that later-breeding cohorts were exposed to in 2018 were more severe in both magnitude and duration than those in 2019. Results In 2018, offspring of the later-breeding cohort were larger but less tolerant of NaCl compared to offspring of the earlier-breeding cohort. The offspring of the earlier-breeding cohort additionally were able to acclimate to a lethal concentration of NaCl following sublethal exposure earlier in life, while the later-breeding cohort became less tolerant of NaCl following acclimation. Interestingly, in 2019, the warmer of the two breeding seasons, we did not detect the negative effects of later breeding phenology on responses to NaCl. Conclusions These results suggest that phenological shifts that expose breeding amphibians to freezing conditions can have cascading consequences on offspring mass and ability to tolerate future stressors but likely depends on the severity of the freeze event.
... Current climate model forecasts suggest that events such as air temperatures that increase faster in the spring could be more frequent with global warming (Príncipe et al. 2017), and theoretically, late spring frost damage occurs more severely as a result of the early opening of the buds in such conditions (Delpierre et al. 2016). In the last 15 years, indeed, it has been reported that late spring frost events cause significant yield losses in different countries, including the USA in 2012 (Kistner et al. 2018;Hufkens et al. 2012) and 2007 (Augspurger 2009), and France in 1995 (Ningre and Colin 2007); and southern Germany, Spain, Switzerland, and northeastern France in 2016 and 2011 (Vitasse and Rebetez 2018), after the onset of early vegetation in the spring. An unusually warm period in these regions has caused bud burst earlier than normal time. ...
Article
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To boost our understanding of recent frost damage events in apricot trees, we focused on estimating the cell death point in the receptacle and pistil organs of apricots by simulating the unexpected temperature changes occurring in early spring under laboratory-based freezing experiments. The hypothesis also that organic acids (oxalic, propionic, tartaric, butyric, malonic, malic, lactic, citric, maleic, fumaric and succinic acids) and soluble sugar compositions (glucose, fructose, sucrose and total sugar) may directly stimulate the change of the frost tolerance in apricot receptacle and pistil organs was investigated using two apricot (Prunus armeniaca L.) cultivars, one tolerant to frost (‘Iğdır Şalak’) and the other sensitive to frost (‘Mihralibey’). Our results indicated that the mean the cell death point (CDP) of flower pistil (from −13.26 to −14.18 °C) was at lower temperatures than those of flower receptacle (from −6.28 to −8.65 °C) in both apricot cultivars during the full blooming stage. In both apricot cultivars, receptacle organs showed more sensitive changes to the frost tolerance in response to low temperatures compared with pistil organs. In terms of organic acid and soluble sugar contents, significant differences were determined for both apricot cultivars and between the pistil and the receptacle organs of the flowers. Organic acid content was higher in the freezing-tolerant apricot cultivar (‘Iğdır Şalak’) than the freezing-sensitive apricot cultivar (‘Mihralibey’), but it was lower in the freezing-tolerant organ (pistil) in comparison with the freezing-sensitive organ (receptacle). Moreover, fructose concentrations for both ‘Mihralibey’ and ‘Iğdır Şalak’ were significantly higher in the receptacle compared to the pistil. A significant negative correlation was also observed between the mean CDP values and both all organic acids and sucrose contents in the pistil and the receptacle organs for both apricot cultivars. There was a positive relationship between the CDP values and fructose content in ‘Iğdır Şalak’. Additionally, there was a negative correlation between the CDP values and sucrose in 'Mihralibey'. Overall, the data presented suggest that the high level of sucrose and organic acid content of receptacle and pistil organs contributes to support frost tolerance of organs. The results of this study could help in the understanding of how receptacle and pistil organs of two different apricot cultivars react to frost stress, and how they modulate their soluble sugar and organic acid metabolism.
... Sugar maple is generally healthy across its range, though declining growth and crown dieback have been noted in selected stands in the last 40 years (Bauce and Allen 1992;Payette et al. 1996). The specific drivers of decline, often termed sugar maple decline disease, are diverse and often locally specific but well studied (Horsley et al. 2002;Hufkens et al. 2012). Given the species' specific environmental (Sullivan et al. 2013;Bauce and Allen 1992;Duchesne et al. 2002) and climate (Gavin 2008) tolerances, climate change is likely to be the largest threat to the species in the future (Bishop et al. 2015). ...
Article
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Given the large contribution of forests to terrestrial carbon storage, there is a need to resolve the environmental and physiological drivers of tree-level response to rising atmospheric CO 2 . This study examines how site-level soil moisture influences growth and intrinsic water-use efficiency in sugar maple (Acer saccharum Marsh.). We construct tree-ring, δ ¹⁸ O, and Δ ¹³ C chronologies for trees across a soil moisture gradient in Ontario, Canada, and employ a structural equation modelling approach to ascertain their climatic, ontogenetic, and environmental drivers. Our results support previous evidence for the presence of strong developmental effects in tree-ring isotopic chronologies — in the range of −4.7‰ for Δ ¹³ C and +0.8‰ for δ ¹⁸ O — across the tree life span. Additionally, we show that the physiological response of sugar maple to increasing atmospheric CO 2 depends on site-level soil moisture variability, with trees only in relatively wet plots exhibiting temporal increases in intrinsic water-use efficiency. These results suggest that trees in wet and mesic plots have experienced temporal increases in stomatal conductance and photosynthetic capacity, whereas trees in dry plots have experienced decreases in photosynthetic capacity. This study is the first to examine sugar maple physiology using a dendroisotopic approach and broadens our understanding of carbon–water interactions in temperate forests.
... Whether APV-induced spring vegetation variability has negative or positive seasonal compensation effects in the subsequent summer and autumn is another important issue to address. Previous studies have shown that the spring environmental stresses, such as low temperatures and drought lasting several months, can greatly reduce vegetation productivity (Noormets et al. 2008;Hufkens et al. 2012). However, Buermann et al. (2018) demonstrated that contrasting lagged productivity responses to spring environmental change exist widely in northern ecosystems, which indicates that spring vegetation variability may not result in the reduction of vegetation productivity. ...
Article
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The Arctic polar vortex (APV) system plays an important role in controlling winter and spring atmospheric circulation pattern in China. Here, we evaluate the anomalous APV-induced spring vegetation variability and lagged productivity responses in China. We found that both strong and weak APV conditions have almost equally negative impacts on spring vegetation growth in China, e.g., negative NDVI anomalies occurred in 48.6 and 53.2% of China’s vegetated areas under strong and weak APV conditions, respectively. However, large seasonal compensation effects were associated with weak APV conditions, and beneficial lagged vegetation productivity responses occurred in 67.2% of China’s vegetated areas, whereas adverse responses occurred in only 32.8% of vegetated areas. Under a strong APV, adverse lagged vegetation productivity responses occurred in 54.5% of China’s vegetated areas, whereas beneficial responses occurred in only 45.5% of vegetated areas. The temperature, precipitation, and solar radiation changes caused by anomalous APV-induced changes in circulation patterns are the main reasons for spring vegetation variability. The lagged vegetation productivity responses were attributed to anomalous APV-related precipitation and air temperature anomalies in the following summer and autumn. This improved understanding of the strong links between APV anomalies and vegetation dynamics in China should facilitate early warning of vegetation productivity reductions under anomalous APV conditions.
... Specifically, a positive relationship of interannual variations of EOS against SOS was found in this study (Fig. 12), which is supported by previous research Liu et al., 2016b). An earlier EOS could result from the risks of spring frost and summer drought that are increased by an earlier SOS (Buermann et al., 2013;Hufkens et al., 2012;Lian et al., 2020). The within-year relationship of a later SOS and EOS maintains a relatively stable growing season length and may reduce the effect of climate warming on terrestrial carbon sequestration . ...
Article
Land surface phenology (LSP) characterizes the seasonal dynamics of vegetation communities that compose individual satellite pixels and its interannual and spatial variations have been widely associated with climate. However, increasing evidence shows an effect of land cover composition within a pixel on LSP, but it remains unclear the extent of impacts relative to other drivers. To fill this gap, this study quantitatively assessed the contributions of land cover composition, climate, and topography on the spatial and interannual variation in LSP throughout the 2002 Ponil Complex Fire in New Mexico, USA, using a machine learning approach of Boosted Regression Trees (BRT). As the fire mainly converted ponderosa pine and Douglas-fir (evergreen tree) to soil ground and Gambel Oak (deciduous shrub), we computed both the proportion of tree cover to all vegetation cover (PTV) and vegetation fractional cover (VFC) as the metrics of land cover composition from high-resolution images in 2018 and from MODIS growing season greenness from 2001-2018. Start (SOS) and end (EOS) of growing season were derived from 500-m MODIS data from 2001-2018 and 30-m Harmonized Landsat Sentinel-2 data in 2018. BRT models showed that PTV was the most important predictor of spatial variations in SOS and EOS in 2018, despite the different contributions (20.3%-42.9%) at 30-m and 500-m spatial scales. Although the growing degree days (28.6%) and the first freeze date (19.6%) were the most important predictors of interannual variations in SOS and EOS from 2001-2018, respectively, VFC also presented an important contribution for SOS (8.4%) and EOS (12.2%). This study demonstrates the utility of machine learning in modeling phenology and highlights the essential role of land cover composition in understanding the spatial and interannual variations of LSP that have been widely associated with topography and climate.
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Phenology is a key indicator of climate change. We studied the phenological change of Populus euphratica during 1960–2019, and investigated the relationships between phenological change and climate change. The results showed that the annual average temperature in the Lower reaches of Heihe River was increased significantly over the past 58years, which directly led to the germination of Populus euphratica was advanced, and the leaf yellowing delayed. The growth season of Populus euphratica was significantly prolonged after 1990, and the mean growth season length were 230, 236, 245 days in 1960–1969,1990–1999,2010–2019, respectively. Temperature is the major environmental factor affecting phenological change, and the length of growing season was prolonged with the increase of annual average temperature and accumulated temperature. Phenological changes have an important effect on plant growth. Our study showed that the frost days of Populus euphratica increasing from 1990 to 2019, and the frost days was increased with the growth season length. The accumulated frost days were 19, 21, 23 days in 1990–1999, 2000–2009, 2009–2019. Our studies also indicated that phenological changes also affect socioeconomic, especially the delay of leaf yellow period maybe has an important impact on tourism due to the best viewing period is postponed. Our study of phenological change law and its relationships with climate changes could guide the government formulates tourism policies and help tourists to arrange best viewing time.
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This chapter assesses climate information relevant to regional impact and risk assessment. It complements other WG1 chapters which focus on the physical processes determining changes in the climate system and on methods for estimating regional changes. This chapter is new in the IPCC WGI assessment reports, in that it represents a contribution to the “IPCC Risk Framework”. Within this framework, climate-related impacts and risks are determined through an interplay between the occurrence of climate hazards and their consequences depending on the exposure of the affected human or natural system and its vulnerability to the hazardous conditions. In Chapter 12, we are assessing climatic impact-drivers that could lead to hazards or to opportunities, from the literature and model results since AR5. This will particularly support the assessment of key risks related to climate change by WGII (Chapter 16). Despite the fact that impacts may also be induced by climate adaptation and mitigation policies themselves, as well as by socioeconomic trends, changes in vulnerability or exposure, and external geophysical hazards such as volcanoes, the focus here is only on ‘climatic’ impacts and risks induced by shifts in physical climate phenomena that directly influence human and ecological systems.
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Changes in climate are leading to modifications in the timing of seasonal events such as migrations and flowering. Erythronium americanum (trout lily) can break bud early in response to warming, but changes to its growing season may be limited by early shade from canopy trees and frost. I experimentally assessed the impact of shade and frost on senescence in E. americanum and descriptively monitored the response of E. americanum to vernal air and soil temperatures in a garden setting. Early shade did not affect the timing of senescence. Experimental exposure to frost resulted in increased leaf damage, earlier senescence, and greater corm death than in control plants. Despite ten days in which the air temperature dropped below freezing, there was no evidence of leaf damage in the field. These results suggest that early shade from canopy trees will not hasten the end of the future growing season for E. americanum, but that late frost could bring about early senescence if that frost is sufficiently hard.
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Climate change is having many effects in the agricultural sector, which are being studied worldwide. Undoubtedly, warmer winters and earlier springs produce changes in frost regimes and severity that will affect the sustainability of agricultural production in the area. The Mediterranean region and the Iberian Peninsula (IP) are among the areas where the greatest impact of climate change is expected. Daily data from 68 weather stations of the IP belonging to the European Climate Assessment and Dataset (1975–2018) were used to conduct a spatiotemporal study of the frost regime. The variables calculated include the probability of three frost types according to their severity, frost day, mean absolute minimum yearly temperature, first frost day, last frost day, and frost-free period. These variables were integrated into a geographic information system, which allowed the graphical visualization of their patterns using of geostatistical interpolation techniques (kriging). Changes in frost variables were investigated using the Mann–Kendall test and Sen’s slope estimator. A general reduction in the number of frosts per year is observed (values between −0.04- and −0.8-day frosts per year), as well as an increase in the mean absolute minimum temperature (values between 0.04 and 0.10 °C per year), with very high significant trends throughout the territory. The reduction in the number of frosts is more pronounced at a higher elevation. Frost dates vary greatly due to the orographic characteristics of the IP. The generalized trend is of a significant delay of the autumn frosts (values between 0.4 and 1.06 days/year), as well as early spring frosts (between −0.429 and −1.29 days/year), and as a consequence a longer frost-free period, all changes were much stronger than those found in other regions of the world. These effects of climate change must be mitigated by modifying species, varieties, and cultivation techniques to guarantee sustainable agriculture.
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Spring phenology is advancing with warming but late spring freezes may not advance at the same rate, potentially leading to an increase in freezes after trees initiate budburst. Research suggests warming winters may delay budburst through reduced chilling, which may cause plants to leaf out more slowly, thus decreasing spring freeze tolerance. Here, we assessed the effects of late spring freezes and reduced over-winter chilling on sapling phenology, growth and tissue traits, across eight temperate tree and shrub species in a laboratory experiment. We found that spring freezes delayed leafout-extending the period of greatest risk for freeze damage-and increased damage to the shoot apical meristem, decreased leaf toughness and leaf thickness. Longer chilling accelerated budburst and leafout, even under spring freeze conditions. Thus chilling compensated for the adverse effects of late spring freezes on phenology. Despite the effects of spring freezes and chilling on phenology, we did not see any major re-ordering in the sequence of species leafout. Our results suggest climate change may impact forest communities not through temporal reassembly, but rather through impacts on phenology and growth from the coupled effects of late spring freezes and decreased over-winter chilling under climate change.
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At the time of leaf emergence in deciduous forests, markedly enhanced evapotranspiration leads to a rapid drop in the Bowen ratio. A small fraction of this surface flux alteration converges into the boundary layer, and this can be detected in the mean temperature and humidity daily increments at the surface. A simple technique is presented for identifying this response in surface climate data and extracting time series for the date of spring onset and for the `spring intensity,' a measure of surface energy budget partition change in spring. A tendency Bowen ratio B is found from changes in the daily increment of temperature and humidity in multidecadal averages. The spring date determined using this criterion for stations along the U.S. east coast corresponds to the date at which the normalized difference vegetation index (NDVI) reaches 80% of its seasonal maximum. Northward movement of the vernal front is similar to that obtained using Hopkins' empirical rule; it is linearly related to leaf emergence and flowering dates from the North American lilac phenology network. Spring intensity increases northward; the states from Virginia north exhibit distinctly higher values. There has been a trend in the most recent decades toward earlier spring dates, except for regions in Virginia and North Carolina. The same analyses performed using the small subset of stations with longer-term records indicate that a trend toward an earlier spring date is confined to recent decades. An inverse relationship between the spring date and spring average temperature was found for the Midwest but is inadequate for the northeast. Spring intensity has generally increased in northeastern North America throughout the twentieth century. However, large oscillations with an approximate 20-yr period distinguish the northeastern United States from the Midwest, indicating that the intensity of spring is not a simple function of spring temperature or of forest cover fraction.
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Projected climate warming will potentially have profound effects on the earth's biota, including a large redistribution of tree species. We developed models to evaluate potential shifts for 80 individual tree species in the eastern United States. First, environmental factors associated with current ranges of tree species were assessed using geographic information systems (GIS) in conjunction with regression tree analysis (RTA). The method was then extended to better understand the potential of species to survive and/ or migrate under a changed climate. We collected, summarized, and analyzed data for climate, soils, land use, elevation, and species assemblages for .2100 counties east of the 100th meridian. Forest Inventory Analysis (FIA) data for .100 000 forested plots in the East provided the tree species range and abundance information for the trees. RTA was used to devise prediction rules from current species-environment relationships, which were then used to replicate the current distribution as well as predict the future potential distri- butions under two scenarios of climate change with twofold increases in the level of at- mospheric CO2. Validation measures prove the utility of the RTA modeling approach for mapping current tree importance values across large areas, leading to increased confidence in the predictions of potential future species distributions. With our analysis of potential effects, we show that roughly 30 species could expand their range and/or weighted importance at least 10%, while an additional 30 species could decrease by at least 10%, following equilibrium after a changed climate. Depending on the global change scenario used, 4-9 species would potentially move out of the United States to the north. Nearly half of the species assessed (36 out of 80) showed the potential for the ecological optima to shift at least 100 km to the north, including seven that could move .250 km. Given these potential future distributions, actual species redistributions will be controlled by migration rates possible through fragmented landscapes.
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In the last two decades the availability of global remote sensing data sets has provided a new means of studying global patterns and dynamics in vegetation. The vast majority of previous work in this domain has used data from the Advanced Very High Resolution Radiometer, which until recently was the primary source of global land remote sensing data. In recent years, however, a number of new remote sensing data sources have become available that have significantly improved the capability of remote sensing to monitor global ecosystem dynamics. In this paper, we describe recent results using data from NASA's Moderate Resolution Imaging Spectroradiometer to study global vegetation phenology. Using a novel new method based on fitting piecewise logistic models to time series data from MODIS, key transition dates in the annual cycle(s) of vegetation growth can be estimated in an ecologically realistic fashion. Using this method we have produced global maps of seven phenological metrics at 1-km spatial resolution for all ecosystems exhibiting identifiable annual phenologies. These metrics include the date of year for (1) the onset of greenness increase (greenup), (2) the onset of greenness maximum (maturity), (3) the onset of greenness decrease (senescence), and (4) the onset of greenness minimum (dormancy). The three remaining metrics are the growing season minimum, maximum, and summation of the enhanced vegetation index derived from MODIS. Comparison of vegetation phenology retrieved from MODIS with in situ measurements shows that these metrics provide realistic estimates of the four transition dates identified above. More generally, the spatial distribution of phenological metrics estimated from MODIS data is qualitatively realistic, and exhibits strong correspondence with temperature patterns in mid- and high-latitude climates, with rainfall seasonality in seasonally dry climates, and with cropping patterns in agricultural areas.
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In this study we seek empirical relationships among canopy resistance to water vapor transport, the time-varying leaf area index (LAI), in situ radiative flux observations, and a satellite-based estimate of leaf state (NDVI, the normalized difference vegetation index) from a leafless deciduous forest to a covered canopy and vice versa. These relationships can be used in numerical models such as verification in global climate models. They also can be useful tools for developing remote sensing techniques.LAI was found through analysis of frequent video images of canopy evolution in spring and autumn during 1992 and 1993 at a deciduous forest in central Massachusetts. We examined the impact of leaf presence on water vapor transport during spring and autumn using an LAI time series during leaf emergence and leaf fall for the four study seasons. The canopy resistance to water vapor transport (rc) decreased abruptly at leaf emergence in each year but then also continued to decrease slowly during the remaining growing season, owing to slowly increasing LAI. One remarkable result is that a single linear relationship between rc. and LAI during leaf emergence can be used to estimate the minimum seasonal rc associated with the maximum foliage cover. Canopy resistance and PAR-albedo (albedo from photosynthetically active radiation (PAR) instruments) began to increase about 1 month before leaf fall with the diminishment of CO2 gradient above the canopy as well, at which time evaporation began to be controlled as if the canopy were leafless. We present empirical linear regressions relating NDVI, rc, and PAR-albedo. The NDVI linear regressions with surface measurements indicate that tower-based measurements can represent at least a satellite pixel region. These results reinforce the notion that relationships among these parameters are scale independent from tower-based measurements spatial scale to a satellite pixel resolution (1.1 km X 1.1 km area)., at least.
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Studies of the effects of climate change on forests have focused on the ability of species to tolerate temperature and moisture changes and to disperse, but they have ignored the effects of disturbances caused by climate change (e.g., Ojima et al. 1991).Yet modeling studies indicate the importance of climate effects on disturbance regimes (He et al. 1999). Local, regional, and global changes in temperature and precipitation can influence the occurrence, timing, frequency, duration, extent, and intensity of disturbances (Baker 1995, Turner et al. 1998). Because trees can survive from decades to centuries and take years to become established, climate-change impacts are expressed in forests, in part, through alterations in disturbance regimes (Franklin et al. 1992, Dale et al. 2000). yes
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A causal relationship between Holocene patterns of plant distributions in the American Southwest and shifts in atmospheric flow structure has been inferred, but has yet to be demonstrated. To investigate this inferred relationship, seedlings of Quercus gambelii were transplanted along various local microhabitat gradients and regional air mass gradients and assessed for patterns of seedling establishment. The primary cause of seedling mortality appeared to be summer drought stress, which paralleled the latitudinal gradient of the `Arizona monsoon'. The `Arizona monsoon' air mass gradient produces an increasing probability of summer drought stress with increasing latitude, while the `polar front' air mass gradient produces an increasing probability of winter and spring cold stress with increasing latitude. We hypothesize that the two gradients produce, as a function of increasing latitude, convergent upper and lower elevational limits of Q. gambelii. The results of this elevational convergence appear to be a virtual absence of oak seedling establishment near the species' northern limits and a gradual senescence of long-lived clones from spring freeze stress when followed by summer drought stress. These conclusions have been extended to the biogeography of Q. turbinella through growth chamber experiments on the comparative drought physiology of seedlings of the two species and through previous work on their associated biogeographic patterns. Thus, the northern ecotones of Q. gambelii and Q. turbinella, appear to be primarily caused by the combination of spring freeze stress and summer moisture stress, rather than freeze stress alone, as previously proposed. We report systematic temporal and spatial shifts in the two air mass gradients, between warming and cooling trends, which can account for the Holocene biogeographic history of the two oak species and, in part, the observed spatial and temporal variations in oak morphology and oak/pine community structure.
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(1) The dates of budhurst of lateral shoots on 2- to 10-year old trees of Picea sitchensis were recorded on fourteen occasions at sites near meteorological stations in lowland and upland Britain between 1960 and 1980. (2) The following relationship accounted for 92% of the variation in thermal time from 1 February to the date of budburst among the fourteen observations: thermal time = 67.4 + 4401.8 exp (-0.042 x chill days) where thermal time was day degrees >5 ⚬C accumulated from 1 February, and chill days were the number of days ⩽5 ⚬C counted from 1 November, both based on mean daily air temperature ((max. + min.)/2). This model may be used to estimate the date of budburst on young P. sitchensis of most provenances growing in upland Britain. (3) The following features or assumptions of the model were examined with reference to the literature and/or by experimentation: the small effect of provenance; linearity in the relationship between bud growth rate and temperature; the large effect of chilling on thermal time to budhurst; the omission of daylength and soil temperature as variables; the choice of starting dates for effective chilling and thermal time; and the use of simple fixed base temperatures. (4) The model was applied to mean daily temperatures at Eskdalemuir for the period 1912-82. The predicted dates of budburst ranged from 23 April in 1961 to 30 May in 1923, with a mean date of 12 May.
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PREFACE TO THE SECOND EDITION LIST OF SYMBOLS 1. SCOPE OF ENVIRONMENTAL PHYSICS 2. GAS LAWS Pressure, volume and temperature Specific heats Lapse rate Water and water vapour Other gases 3. TRANSPORT LAWS General transfer equation Molecular transfer processes Diffusion coefficients Radiation laws 4. RADIATION ENVIRONMENT Solar radiation Terrestrial radiation Net radiation 5. MICROCLIMATOLOGY OF RADIATION (i) Interception Direct solar radiation Diffuse radiation Radiation in crop canopies 6. MICROCLIMATOLOGY OF RADIATION (ii) Absorption and reflection Radiative properties of natural materials Net radiation 7. MOMENTUM TRANSFER Boundary layers Wind profiles and drag on uniform surfaces Lodging and windthrow 8. HEAT TRANSFER Convection Non-dimensional groups Measurements of convection Conduction Insulation of animals 9. MASS TRANSFER (i) Gases and water vapour Non-dimensional groups Measurement of mass transfer Ventilation Mass transfer through pores Coats and clothing 10.MASS TRANSFER (ii) Particles Steady motion 11.STEADY STATE HEAT BALANCE (i) Water surfaces and vegetation Heat balance equation Heat balance of thermometers Heat balance of surfaces Developments from the Penman Equation 12.STEADY STATE HEAT BALANCE (ii) Animals Heat balance components The thermo-neutral diagram Specification of the environment Case studies 13.TRANSIENT HEAT BALANCE Time constant General cases Heat flow in soil 14.CROP MICROMETEOROLOGY (i) Profiles and fluxes Profiles Profile equations and stability Measurement of flux above the canopy 15.CROP MICROMETEOROLOGY (ii) Interpretation of measurements Resistance analogues Case studies: Water vapour and transpiration Carbon dioxide and growth Sulphur dioxide and pollutant fluxes to crops Transport within canopies APPENDIX BIBLIOGRAPHY REFERENCES INDEX