Accurate phenology analyses require bud traits and energy budgets.

Spring phenology is mainly driven by temperature in extratropical ecosystems. Recent evidence highlighted the key role of micrometeorology and bud temperature on delaying or advancing leaf unfolding. Yet, phenology studies, either using ground-based or remote sensing observations, always substitute plant tissue temperature by air temperature. In fact, temperatures differ substantially between plant tissues and the air because plants absorb and lose energy. Here, we build on recent observations and well-established energy balance theories to discuss how solar radiation, wind and bud traits might affect our interpretation of spring phenology sensitivity to warming. We show that air temperature might be an imprecise and biased predictor of bud temperature. Better characterizing the plants' phenological response to warming will require new observations of bud traits and temperature for accurately quantifying their energy budget. As consistent micrometeorology datasets are still scarce, new approaches coupling energy budget modelling and plant traits could help to improve phenology analyses across scales.

High photosynthetic capacity of Sahelian C3 and C4 plants.

The semi-arid ecosystems of the African Sahel play an important role in the global carbon cycle and are among the most sensitive ecosystems to future environmental pressures. Still, basic data of photosynthetic characteristics of Sahelian vegetation are very limited, preventing us to properly understand these ecosystems and to project their response to future global changes. Here, we aim to study and quantify key leaf traits, including photosynthetic parameters and leaf nutrients (Nleaf and Pleaf), of common C3 and C4 Sahelian plants (trees, lianas, and grasses) at the Dahra field site (Senegal). Dahra is a reference site for grazed semi-arid Sahelian savannah ecosystems in carbon cycle studies. Within the studied species, we found that photosynthetic parameters varied considerably between functional types. We also found significant relationships between and within photosynthetic parameters and leaf traits which mostly differed in their slopes from C3 to C4 plants. In agreement with the leaf economic spectrum, strong relationships (R2 = 0.71) were found between SLA and Nleaf whereby C3 and C4 plants showed very similar relationships. By comparing our data to a global dataset of plant traits, we show that measured Sahelian plants exhibit higher photosynthetic capacity (Asat) compared to the non-Sahelian vegetation, with values that are on average a fourfold of the global average. Moreover, Sahelian C3 plants showed photosynthetic nutrient use efficiencies that were on average roughly twice as high as global averages. We interpreted these results as the potential adaptation of Sahelian plants to short growing season lengths via an efficient nutrient allocation to optimize photosynthesis during this period. Our study provides robust estimates of key functional traits, but also traits relationships that will help to calibrate and validate vegetation models over this data-poor region.

Soil thawing regulates the spring growth onset in tundra and alpine biomes.

Soil temperature remains isothermal at 0 °C and water shifts to a liquid phase during soil thawing. Vegetation may receive this process as a signal and a key to restore physiological activity. We aimed to show the relationship between the timing of soil thawing and the spring growth onset. We estimated the delay between the soil thawing and the spring growth onset in 78 sites of the FLUXNET network. We built a soil thawing map derived from modeling for the northern hemisphere and related it to the greenness onset estimated with satellite imagery. Spring onset estimated with GPP time series occurred shortly after soil surface thawing in tundra (1.1 ± 3.5 days) and alpine grasslands (16.6 ± 5.8 days). The association was weaker for deciduous forests (40.3 ± 4.2 days), especially where soils freeze infrequently. Needleleaved forests tended to start the growing season before the end of thawing (-17.4 ± 3.6 days), although observations from remote sensing (MODIS Land Cover Dynamics) indicated that the onset of greenness started after the thawing period (26.8 ± 3.2 days). This study highlights the role of soil temperature at the spring growth onset at high latitudes. Soil thawing becomes less relevant in temperate forests, where soil is occasionally frozen and other climate factors become more important.

Spatial variance of spring phenology in temperate deciduous forests is constrained by background climatic conditions.

Leaf unfolding in temperate forests is driven by spring temperature, but little is known about the spatial variance of that temperature dependency. Here we use in situ leaf unfolding observations for eight deciduous tree species to show that the two factors that control chilling (number of cold days) and heat requirement (growing degree days at leaf unfolding, GDDreq) only explain 30% of the spatial variance of leaf unfolding. Radiation and aridity differences among sites together explain 10% of the spatial variance of leaf unfolding date, and 40% of the variation in GDDreq. Radiation intensity is positively correlated with GDDreq and aridity is negatively correlated with GDDreq spatial variance. These results suggest that leaf unfolding of temperate deciduous trees is adapted to local mean climate, including water and light availability, through altered sensitivity to spring temperature. Such adaptation of heat requirement to background climate would imply that models using constant temperature response are inherently inaccurate at local scale.

Velocity of change in vegetation productivity over northern high latitudes.

Warming is projected to increase the productivity of northern ecosystems. However, knowledge on whether the northward displacement of vegetation productivity isolines matches that of temperature isolines is still limited. Here we compared changes in the spatial patterns of vegetation productivity and temperature using the velocity of change concept, which expresses these two variables in the same unit of displacement per time. We show that across northern regions (>50° N), the average velocity of change in growing-season normalized difference vegetation index (NDVIGS, an indicator of vegetation productivity; 2.8 ± 1.1 km yr-1) is lower than that of growing-season mean temperature (T GS; 5.4 ± 1.0 km yr-1). In fact, the NDVIGS velocity was less than half of the T GS velocity in more than half of the study area, indicating that the northward movement of productivity isolines is much slower than that of temperature isolines across the majority of northern regions (about 80% of the area showed faster changes in temperature than productivity isolines). We tentatively attribute this mismatch between the velocities of productivity and temperature to the effects of limited resource availability and vegetation acclimation mechanisms. Analyses of ecosystem model simulations further suggested that limited nitrogen availability is a crucial obstacle for vegetation to track the warming trend.

Three times greater weight of daytime than of night-time temperature on leaf unfolding phenology in temperate trees.

The phenology of spring leaf unfolding plays a key role in the structure and functioning of ecosystems. The classical concept of heat requirement (growing degree days) for leaf unfolding was developed hundreds of years ago, but this model does not include the recently reported greater importance of daytime than night-time temperature. A manipulative experiment on daytime vs night-time warming with saplings of three species of temperate deciduous trees was conducted and a Bayesian method was applied to explore the different effects of daytime and night-time temperatures on spring phenology. We found that both daytime and night-time warming significantly advanced leaf unfolding, but the sensitivities to increased daytime and night-time temperatures differed significantly. Trees were most sensitive to daytime warming (7.4 ± 0.9, 4.8 ± 0.3 and 4.8 ± 0.2 d advancement per degree Celsius warming (d °C-1 ) for birch, oak and beech, respectively) and least sensitive to night-time warming (5.5 ± 0.9, 3.3 ± 0.3 and 2.1 ± 0.9 d °C-1 ). Interestingly, a Bayesian analysis found that the impact of daytime temperature on leaf unfolding was approximately three times higher than that of night-time temperatures. Night-time global temperature is increasing faster than daytime temperature, so model projections of future spring phenology should incorporate the effects of these different temperatures.

Declining global warming effects on the phenology of spring leaf unfolding.

Earlier spring leaf unfolding is a frequently observed response of plants to climate warming. Many deciduous tree species require chilling for dormancy release, and warming-related reductions in chilling may counteract the advance of leaf unfolding in response to warming. Empirical evidence for this, however, is limited to saplings or twigs in climate-controlled chambers. Using long-term in situ observations of leaf unfolding for seven dominant European tree species at 1,245 sites, here we show that the apparent response of leaf unfolding to climate warming (ST, expressed in days advance of leaf unfolding per °C warming) has significantly decreased from 1980 to 2013 in all monitored tree species. Averaged across all species and sites, ST decreased by 40% from 4.0 ± 1.8 days °C(-1) during 1980-1994 to 2.3 ± 1.6 days °C(-1) during 1999-2013. The declining ST was also simulated by chilling-based phenology models, albeit with a weaker decline (24-30%) than observed in situ. The reduction in ST is likely to be partly attributable to reduced chilling. Nonetheless, other mechanisms may also have a role, such as 'photoperiod limitation' mechanisms that may become ultimately limiting when leaf unfolding dates occur too early in the season. Our results provide empirical evidence for a declining ST, but also suggest that the predicted strong winter warming in the future may further reduce ST and therefore result in a slowdown in the advance of tree spring phenology.