A modelling analysis of the effects of climatic warming on the probability of spring frost damage to tree species in The Netherlands and Germany

Two studies presented in the literature ( Murray, Canned & Smith 1989 ; Hanninen 1991 ) evaluate the effect of increasing winter temperature on the probability of spring frost damage to trees, but yield contradictory results. It is unclear whether the disparity can be ascribed to the fact that different models were used, or is the result of different climatic warming scenarios being used, or is because the tree species at the different locations do indeed respond differently to warmer winters. To evaluate the effects of climatic warming to tree species in The Netherlands and in Germany, both models were fitted to long series of observations on the date of leaf unfolding of eleven tree species. The impact of the two scenarios (uniformly and non-uniformly changing winter temperature) on the date of leaf unfolding and on the probability of freezing temperature around that date was evaluated. To test the importance of adaptation to local climate, hypothetical provenance transfers were analysed. It was concluded that, for tree species in The Netherlands and Germany, the probability of spring frost damage will decrease. The contradictory results found in the literature could be ascribed to differences between provenances adapted to their local climate, and is not because different models and different climatic warming scenarios were used in these studies.     

Long-term temporal changes of plant phenology in the Western Mediterranean

Plants are altering their life cycles in response to current climatic change around the globe. More than 200 000 records for six phenological events (leaf unfolding, flowering, fruit ripening, fruit harvesting, leaf falling and growing season) of 29 perennial species for the period 1943–2003 recorded throughout Spain provide the longest temporal and the broadest spatial assessment of plant phenology changes in the Mediterranean region. The overwhelming majority of the 118 studied phenophases shifted their dates in recent decades. Such changes differed among phenological events. Leaf unfolding, flowering and fruiting are markedly advancing (?0.48, ?0.59 and ?0.32 days yr^?1, respectively), but only since the mid‐1970s. Anemophilous have advanced more days their flowering than entomophilous. However, some species have delayed and others have advanced their leaf falling dates and as a result only a weak shift was observed in this event for the whole of the studied species (+0.12 days yr^?1). The growing season lengthened by 18 days, which implies an increase of 8% in the life of annual leaves. Such an increase was achieved mainly through the advance of leaf unfolding dates in the spring, one of the most productive times of year for vegetation in the Mediterranean. Shifts in the plant calendar were accompanied as well by long‐term changes in the range of onset dates in 39% of studied phenophases. Leaf unfolding, flowering and growing season tended to reduce spatial variability, reflecting a faster and more synchronized onset (or duration) of phenophases across the study area. Changes in spatial variability may aggravate calendar mismatching with other trophic levels resulting from changes in dates. Because temporal responses differed markedly among species, calendar guilds of plants have changed, which suggests alterations of interspecific relationships in plant communities from Mediterranean ecosystems.     

Impact of climate change on plant phenology in Mediterranean ecosystems

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

Responses of canopy duration to temperature changes in four temperate tree species: relative contributions of spring and autumn leaf phenology

While changes in spring phenological events due to global warming have been widely documented, changes in autumn phenology, and therefore in growing season length, are less studied and poorly understood. However, it may be helpful to assess the potential lengthening of the growing season under climate warming in order to determine its further impact on forest productivity and C balance. The present study aimed to: (1) characterise the sensitivity of leaf phenological events to temperature, and (2) quantify the relative contributions of leaf unfolding and senescence to the extension of canopy duration with increasing temperature, in four deciduous tree species (Acer pseudoplatanus, Fagus sylvatica, Fraxinus excelsior and Quercus petraea). For 3 consecutive years, we monitored the spring and autumn phenology of 41 populations at elevations ranging from 100 to 1,600 m. Overall, we found significant altitudinal trends in leaf phenology and species-specific differences in temperature sensitivity. With increasing temperature, we recorded an advance in flushing from 1.9 ± 0.3 to 6.6 ± 0.4 days °C⁻¹ (mean ± SD) and a 0 to 5.6 ± 0.6 days °C⁻¹ delay in leaf senescence. Together both changes resulted in a 6.9 ± 1.0 to 13.0 ± 0.7 days °C⁻¹ lengthening of canopy duration depending on species. For three of the four studied species, advances in flushing were the main factor responsible for lengthening canopy duration with increasing temperature, leading to a potentially larger gain in solar radiation than delays in leaf senescence. In contrast, for beech, we found a higher sensitivity to temperature in leaf senescence than in flushing, resulting in an equivalent contribution in solar radiation gain. These results suggest that climate warming will alter the C uptake period and forest productivity by lengthening canopy duration. Moreover, the between-species differences in phenological responses to temperature evidenced here could affect biotic interactions under climate warming.     

European phenological response to climate change matches the warming pattern

Global climate change impacts can already be tracked in many physical and biological systems; in particular, terrestrial ecosystems provide a consistent picture of observed changes. One of the preferred indicators is phenology, the science of natural recurring events, as their recorded dates provide a high-temporal resolution of ongoing changes. Thus, numerous analyses have demonstrated an earlier onset of spring events for mid and higher latitudes and a lengthening of the growing season. However, published single-site or single-species studies are particularly open to suspicion of being biased towards predominantly reporting climate change-induced impacts. No comprehensive study or meta-analysis has so far examined the possible lack of evidence for changes or shifts at sites where no temperature change is observed. We used an enormous systematic phenological network data set of more than 125 000 observational series of 542 plant and 19 animal species in 21 European countries (1971–2000). Our results showed that 78% of all leafing, flowering and fruiting records advanced (30% significantly) and only 3% were significantly delayed, whereas the signal of leaf colouring/fall is ambiguous. We conclude that previously published results of phenological changes were not biased by reporting or publication predisposition: the average advance of spring/summer was 2.5 days decade⁻¹ in Europe. Our analysis of 254 mean national time series undoubtedly demonstrates that species' phenology is responsive to temperature of the preceding months (mean advance of spring/summer by 2.5 days°C⁻¹, delay of leaf colouring and fall by 1.0 day°C⁻¹). The pattern of observed change in spring efficiently matches measured national warming across 19 European countries (correlation coefficient r =−0.69, P <0.001).     

Olive phenology as a sensitive indicator of future climatic warming in the Mediterranean

Experimental and modelling work suggests a strong dependence of olive flowering date on spring temperatures. Since airborne pollen concentrations reflect the flowering phenology of olive populations within a radius of 50 km, they may be a sensitive regional indicator of climatic warming. We assessed this potential sensitivity with phenology models fitted to flowering dates inferred from maximum airborne pollen data. Of four models tested, a thermal time model gave the best fit for Montpellier, France, and was the most effective at the regional scale, providing reasonable predictions for 10 sites in the western Mediterranean. This model was forced with replicated future temperature simulations for the western Mediterranean from a coupled ocean‐atmosphere general circulation model (GCM). The GCM temperatures rose by 4·5 °C between 1990 and 2099 with a 1% per year increase in greenhouse gases, and modelled flowering date advanced at a rate of 6·2 d per °C. The results indicated that this long‐term regional trend in phenology might be statistically significant as early as 2030, but with marked spatial variation in magnitude, with the calculated flowering date between the 1990s and 2030s advancing by 3–23 d. Future monitoring of airborne olive pollen may therefore provide an early biological indicator of climatic warming in the Mediterranean.     

Phenotypic plasticity in response to light in the coffee tree

Phenotypic plasticity to light availability was examined at the leaf level in field-grown coffee trees (Coffea arabica). This species has been traditionally considered as shade-demanding, although it performs well without shade and even out-yields shaded coffee. Specifically, we focused our attention on the morpho-anatomical plasticity, the balance between light capture and excess light energy dissipation, as well as on physiological traits associated with carbon gain. A wide natural light gradient, i.e., a diurnal intercepted photon irradiance differing by a factor of 25 between the deepest shade leaves and the more exposed leaves in the canopy, was explored. Responses of most traits to light were non-linear, revealing the classic leaf sun vs. leaf shade dichotomy (e.g., compared with sun leaves, shade leaves had a lower stomatal density, a thinner palisade mesophyll, a higher specific leaf area, an improved light capture, a lower respiration rate, a lower light compensating point and a limited capacity for photoprotection). The light-saturated rates of net photosynthesis were higher in sunlit than in shade leaves, although sun leaves were not efficient enough to use the extra light supply. However, sun leaves showed well-developed photoprotection mechanisms in comparison to shade leaves, which proved sufficient for avoiding photoinhibition. Specifically, a higher non-photochemical quenching coefficient was found in parallel to increases in: (i) zeaxanthin pools, (ii) de-epoxidation state of the xanthophyll cycle, and (iii) activities of some antioxidant enzymes. Intracanopy plasticity depended on the suite of traits considered, and was high for some physiological traits associated with photoprotection and maintenance of a positive carbon balance under low light, but low for most morpho-anatomical features. Our data largely explain the successful cultivation of the coffee tree in both exposed and shade environments, although with a poor resource-use efficiency in high light.     

Leaf phenology in 22 North American tree species during the 21st century

Recent shifts in phenology are the best documented biological response to current anthropogenic climate change, yet remain poorly understood from a functional point of view. Prevailing analyses are phenomenological and approximate, only correlating temperature records to imprecise records of phenological events. To advance our understanding of phenological responses to climate change, we developed, calibrated, and validated process-based models of leaf unfolding for 22 North American tree species. Using daily meteorological data predicted by two scenarios (A2: +3.2 °C and B2: +1 °C) from the HadCM3 GCM, we predicted and compared range-wide shifts of leaf unfolding in the 20th and 21st centuries for each species. Model predictions suggest that climate change will affect leaf phenology in almost all species studied, with an average advancement during the 21st century of 5.0 days in the A2 scenario and 9.2 days in the B2 scenario. Our model also suggests that lack of sufficient chilling temperatures to break bud dormancy will decrease the rate of advancement in leaf unfolding date during the 21st century for many species. Some temperate species may even have years with abnormal budburst due to insufficient chilling. Species fell into two groups based on their sensitivity to climate change: (1) species that consistently had a greater advance in their leaf unfolding date with increasing latitude and (2) species in which the advance in leaf unfolding differed from the center to the northern vs. southern margins of their range. At the interspecific level, we predicted that early-leafing species tended to show a greater advance in leaf unfolding date than late-leafing species; and that species with larger ranges tend to show stronger phenological changes. These predicted changes in phenology have significant implications for the frost susceptibility of species, their interspecific relationships, and their distributional shifts.     

Phenological responses in maple to experimental atmospheric warming and CO2 enrichment

Evidence that global warming has altered the phenology of the biosphere, possibly contributing to increased plant production in the northern hemisphere, has come from a diversity of observations at scales ranging from the view of the back yard to satellite images of the earth. These observations, coupled with an understanding of the effects of temperature on plant phenology, suggest that future changes in the atmosphere and climate could alter plant phenology with unknown or unpredictable consequences. We assessed the effects of simulated climatic warming and atmospheric CO₂ enrichment on the spring and autumn phenology of maple trees (Acer rubrum and A. saccharum) growing for four years in open‐top field chambers. CO₂ enrichment (+300 ppm) had no consistent effects on the timing of budbreak and leaf unfolding in the spring or leaf abscission in the autumn. Warming (+4°C) usually had predictable effects: in two of the three years of assessment, budbreak occurred earlier in warm chambers than in ambient temperature chambers, and leaf abscission always occurred later. The lengthening of the growing season could contribute to increased productivity, although effects of temperature on other physiological processes can concurrently have negative effects on productivity. In 1995, budbreak was unexpectedly delayed in the warmer chambers, apparently the result of advanced budbreak leading to injury from a late‐spring frost. Likewise, there was increased risk associated with longer leaf retention in the autumn: in 1994, leaves in the warm chambers were killed by freezing temperatures before they had senesced. These observations support the premise that global warming could increase the length of the growing season. Phenological responses should, therefore, be part of any assessment of the possible consequences of global change, but our results also suggest that those responses may not always have positive effects on production.     

Changes in autumn senescence in northern hemisphere deciduous trees: a meta-analysis of autumn phenology studies

Background and Aims Many individual studies have shown that the timing of leaf senescence in boreal and temperate deciduous forests in the northern hemisphere is influenced by rising temperatures, but there is limited consensus on the magnitude, direction and spatial extent of this relationship.Methods A meta-analysis was conducted of published studies from the peer-reviewed literature that reported autumn senescence dates for deciduous trees in the northern hemisphere, encompassing 64 publications with observations ranging from 1931 to 2010.Key Results Among the meteorological measurements examined, October temperatures were the strongest predictors of date of senescence, followed by cooling degree-days, latitude, photoperiod and, lastly, total monthly precipitation, although the strength of the relationships differed between high- and low-latitude sites. Autumn leaf senescence has been significantly more delayed at low (25° to 49°N) than high (50° to 70°N) latitudes across the northern hemisphere, with senescence across high-latitude sites more sensitive to the effects of photoperiod and low-latitude sites more sensitive to the effects of temperature. Delays in leaf senescence over time were stronger in North America compared with Europe and Asia.Conclusions The results indicate that leaf senescence has been delayed over time and in response to temperature, although low-latitude sites show significantly stronger delays in senescence over time than high-latitude sites. While temperature alone may be a reasonable predictor of the date of leaf senescence when examining a broad suite of sites, it is important to consider that temperature-induced changes in senescence at high-latitude sites are likely to be constrained by the influence of photoperiod. Ecosystem-level differences in the mechanisms that control the timing of leaf senescence may affect both plant community interactions and ecosystem carbon storage as global temperatures increase over the next century.     

Phenological features in relation to growth forms and biomass accumulation in an alpine meadow of the Central Himalaya

Phenological records of 50 plant species were made during 1988–89 in an alpine meadow (30° 10-30° 13 N lat. and 79° 39-79°-41 E long.) of Central Himalaya located between 3100–3750 m elevation. The growth initiation occurred when temperature began to rise continuously and the resulting in snowmelt. The peaks of the various phenophases succeeded one after another in time, within a period of about four months (from May to September) which is longer than the period reported for the alpine sites of higher latitudes. The period of growth initiation appeared to be related to growth form, the species showing earlier growth initiation (when temperatures were lower) deployed leaves to lower heights, close to the ground. A majority of forbs completed growth cycle earlier than the grasses. For example, one group represented by Trachydium roylei reached peak growth a few weeks before did another group of species, represented by Danthonia cachemyriana, which is indicative of niche separation of the respective group of species. Activities of accumulation of live and dead shoot biomass were clearly separated in time in most communities, the former occurring from May to August, and the latter mainly in September. Nomenclature: Osmaston (1926).     

Flowering phenology in the central highland of Iceland and implications for climatic warming in the Arctic

The cool and short growing season that characterizes Arctic climates puts severe constraints on life cycles and reproduction in the Arctic flora. The timing of flowering is particularly critical and may affect both breeding system and reproductive success through the heavy penalties associated with later flowering. An 11-year study of 75 species in the central highland of Iceland showed that the onset of flowering varies greatly among years. The number of species in flower by the first week of July was closely correlated with air temperature (degree days above zero) in the preceding 5 weeks, but no correlations were found with degree days in May or with total degree days in the previous growing season. Time of snowmelt, which has widely been regarded as the environmental event initiating growth and flowering in alpine and arctic tundra, only had a significant effect when two exceptionally cold and late summers were included. The species studied, most of which have a wide distribution in the Arctic, are predicted to respond quickly to warmer spring and early summer temperatures. Accelerated phenologies may alter patterns of resource allocation, have implications for pollinators and pollinator-competition, and could increase the size, species richness and intraspecific genetic diversity of the soil seed bank. Received: 15 February 1997 / Accepted: 25 October 1997     

Changed plant and animal life cycles from 1952 to 2000 in the Mediterranean region

The available data on climate over the past century indicate that the earth is warming. Important biological effects, including changes of plant and animal life cycle events, have already been reported. However, evidence of such effects is still scarce and has been mostly limited to northern latitudes. Here we provide the first long‐term (1952–2000) evidence of altered life cycles for some of the most abundant Mediterranean plants and birds, and one butterfly species. Average annual temperatures in the study area (Cardedeu, NE Spain) have increased by 1.4 °C over the observation period while precipitation remained unchanged. A conservative linear treatment of the data shows that leaves unfold on average 16 days earlier, leaves fall on average 13 days later, and plants flower on average 6 days earlier than in 1952. Fruiting occurs on average 9 days earlier than in 1974. Butterflies appear 11 days earlier, but spring migratory birds arrive 15 days later than in 1952. The stronger changes both in temperature and in phenophases timing occurred in the last 25 years. There are no significant relationships among changes in phenophases and the average date for each phenophase and species. There are not either significant differences among species with different Raunkiaer life‐forms or different origin (native, exotic or agricultural). However, there is a wide range of phenological alterations among the different species, which may alter their competitive ability, and thus, their ecology and conservation, and the structure and functioning of ecosystems. Moreover, the lengthening of plant growing season in this and other northern hemisphere regions may contribute to a global increase in biospheric activity.     

Phenotypic responses of lacustrine brook charr in relation to the intensity of interspecific competition

We investigated the structure of interindividual variations in the diet of brook charr (Salvelinus fontinalis) based on stomach contents data of 3776 charr captured in 69 lakes of the Canadian Shield (Québec, Canada); 29 of these contained allopatric brook charr populations, 24 contained brook charr and creek chub (Semotilus atromaculatus) and 16 contained brook charr and white sucker (Catostomus commersoni). In any given lake, some of the charr fed almost exclusively on benthic organisms (benthic specialists, i.e., mean percent weight of benthic prey >90%), others, almost exclusively on pelagic prey (pelagic specialists, i.e., mean percent weight of benthic prey <10%), and a lesser proportion were generalist feeders (i.e. mean percent weight of benthic prey between 10 and 90%). The proportion of benthic and pelagic specialists were respectively 41.3 and 18% in allopatric brook charr populations. These proportions fit remarkably well with those based on interindividual preferences in spatial distribution, identified through radio-telemetry in another study done in two lakes of the same area. The proportion of benthic specialists was related to competition for benthic organisms with creek chub and white sucker. The effect of white sucker on brook charr diet was more pronounced than that of creek chub: the proportion of benthic specialists among brook charr decreased from 41.3% in allopatry to 19.7% in sympatry with creek chub, and to 9.9% in sympatry with white sucker. Other response variables of brook charr populations also indicate that white sucker is a stronger competitor than creek chub in this system. Because sucker and chub were introduced in these lakes during the last century, phenotypic responses of brook charr to interspecific competition appear to be rapid. Furthermore, in addition to providing a strong field support to the current hypothesis that polymorphism is promoted by arelaxation of interspecific competition, our results also indicate that phenotypic response of brook charr (i.e. the proportion of each form in a given lake) is related to the intensity of this competition.     

Crown development in a pioneer tree, Rhus trichocarpa, in relation to the structure and growth of individual branches

Based on an allometric reconstruction, the structure and biomass-allocation patterns of branches and current-year shoots were investigated in branches of various heights in the pioneer tree Rhus trichocarpa, to evaluate how crown development is achieved and limited in association with height. Path analysis was conducted to explore the effects of light availability, basal height and size of individual branches on branch structure and growth. Branch angle was affected by basal height, whereas branch mass was influenced primarily by light availability. This result suggests that branch structure is strongly constrained by basal height, and that trees mediate such constraints under different light environments. Previous-year leaf area and light availability showed positive effects on current-year stem mass. In contrast, branch basal height and mass negatively affected current-year stem mass. Moreover, the length of stems of a given diameter decreased with increasing branch height. Therefore the cost of biomass investment for a unit growth in length is greater for branches of larger size and at upper positions. Vertical growth rate in length decreased with increasing height. Height-dependent changes in stem allometry and angle influenced the reduction in vertical growth rate to a similar degree.     

Deciphering the developmental plasticity of walnut saplings in relation to climatic factors and light environment

Developmental plasticity, the acclimation of plants to their local environment, is known to be crucial for the fitness of perennial organisms such as trees. However, deciphering the many possible developmental and environmental influences involved in such plasticity in natural conditions requires dedicated statistical models integrating developmental phases, environmental factors, and interindividual heterogeneity. These models should be able to analyse retrospective data (number of leaves or length of annual shoots along the main stem in the present case). In this study Markov switching linear mixed models were applied to the analysis of the developmental plasticity of walnut saplings during the establishment phase in a mixed Mediterranean forest. In the Markov switching linear mixed models estimated from walnut data sets, the underlying Markov chain represents both the succession and lengths of growth phases, while the linear mixed models represent both the influence of climatic factors and interindividual heterogeneity within each growth phase. On the basis of these integrative statistical models, it is shown that walnut saplings have an opportunistic mode of development that is primarily driven by the changing light environment. In particular, light availability explains the ability of a tree to reach a phase of strong growth where the first branches can appear. It is also shown that growth fluctuation amplitudes in response to climatic factors increased while interindividual heterogeneity decreased during tree development.     

Sociality of Columbian ground squirrels in relation to their seasonal energy intake

Summary Seasonal energy intake was estimated for ten populations of Columbian ground squirrels (Spermophilus columbianus) in northwestern Montana. We calculated daily energy intake for an average ground squirrel in each population using measurements of feeding time, consumption rates of different vegetation types (monocots vs. dicots), and the proportion of monocots and dicots in the diet. These daily energy intakes were multiplied by the length of the plant growing season for each population to estimate seasonal energy intake, i.e. over the ground squirrel active season. Amicable interaction rates measured for each population varied with seasonal energy intake, but not with environmental heterogeneity, sex ratio, or the ratio of adults to juveniles. In particular, amicable interactions among adult-juvenile and juvenile-juvenile pairs increased as seasonal energy intake decreased. The proportion of females breeding as yearlings increased as seasonal energy intake increased. This suggests that harsh environments reduce the energy available for juvenile growth and development, leading to delayed dispersal and age at first reproduction. These responses may promote the formation of kin groups and increased amicable interactions within those groups. The length of the plant growing season may determine environmental harshness across elevational gradients, but at a particular elevation, harshness may depend on factors determining daily food intake.     

Intraspecific variation in spring leaf phenology and duration of leaf expansion in relation to leaf habit and leaf size of temperate tree species

Plant Ecology - Spring leaf phenology has been intensively studied in temperate deciduous broad-leaved tree species, but the phenology of evergreen broad-leaved tree species has seldom been focused...