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Image dataset used for a colour analysis of coral branches throughout a long-term marine heatwave emulation experiment using machine learning. Article: "Within population variability in coral heat tolerance indicates climate adaptation potential" by Humanes and Lachs et al. Code to analyse the dataset is found at 10.5281/zenodo.6256164. LL received funding from Natural Environment Research Council (NERC) ONE Planet Doctoral Training Partnership (NE/S007512/1).

[Experimental design: thermal performance experiments] All experiments were run in climate-controlled incubation facilities of the Institut Mediterrani d’Estudis Avançats (Mallorca, Spain). Following 48 hrs under ambient (collection site) conditions, samples were transferred to individual experimental aquaria, which consisted of a double layered transparent plastic bag filled with 2 L of filtered seawater (60 μm) (following Savva et al. 2018). 16 experimental bags were suspended within 80L temperature-controlled baths. In total, ten baths were used, one for each experimental temperature treatment. Bath temperatures were initially set to the acclimatization temperature (i.e. in situ temperatures) and were subsequently increased or decreased by 1 °C every 24 hours until the desired experimental temperature was achieved. Experimental temperatures were: 15, 18, 21, 24, 26, 28, 30, 32, 34 and 36°C (Table S2). For each species, four replicate aquarium bags were used for each temperature treatment with three individually marked seagrass shoots or three algal fragments placed into each bag. For P. oceanica, each marked plant was a single shoot including leaves, vertical rhizome and roots. For C. nodosa, each marked individual consisted of a 10 cm fragment of horizontal rhizome containing three vertical shoots. Individually marked seaweeds contained the holdfast, and 4-5 fronds of P. pavonica (0.98 ± 0.06 g FW; mean ± SE) or a standardised 5-8 cm fragment with meristematic tip for C. compressa (3.67 ± 0.1 g FW; mean ± SE). Experimental plants were cleaned of conspicuous epiphytes. Once the targeted temperatures were reached in all of the baths, experiments ran for 14 days for the algal species and 21 days for seagrasses to allow for measurable growth in all species at the end of the experiment. Experiments were conducted inside a temperature-controlled chamber at constant humidity and air temperature (15 °C). Bags were arranged in a 4x4 grid within each bath, enabling four species/population treatments to be run simultaneously. Bags were mixed within each bath so that one replicate bag was in each row and column of the grid, to minimise any potential within bath effects of bag position. Replicate bags were suspended with their surface kept open to allow gas exchange and were illuminated with a 14h light:10h dark photoperiod through fluorescent aquarium growth lamps. The water within the bags were mixed with aquaria pumps. The light intensity within each bag was measured via a photometric bulb sensor (LI-COR) and ranged between 180-258 μmol m-2 s-1. Light intensity was constant between experiments and did not significantly differ between experimental treatments (p > 0.05). The temperature in the baths was controlled and recorded with an IKS-AQUASTAR system, which was connected to heaters and thermometers. The seawater within the bags was renewed every 72 hrs and salinity was monitored daily with an YSI multi-parameter meter. Distilled water was added when necessary to ensure salinity levels remained within the range of 36-39 PSU, typical of the study region. Carbon and Nitrogen concentrations in the leaf tissue were measured at the end of the experiment for triplicates of the 24ºC treatment for each species and location (Fig. S2) at Unidade de Técnicas Instrumentais de Análise (University of Coruña, Spain) with an elemental analyser FlashEA112 (ThermoFinnigan). [Species description and distribution] The species used in this study are all common species throughout the Mediterranean Sea, although differ in their biological traits, evolutionary histories and thermo-geographic affinities (Fig. S1). P. oceanica is endemic to the Mediterranean Sea with the all other Posidonia species found in temperate Australia (Aires et al. 2011). The distribution of P. oceanica is restricted to the Mediterranean, spanning from Gibraltar in the west to Cyprus in the east and north into the Aegean and Adriatic seas (Telesca et al. 2015) (Fig. S1A). C. nodosa distribution extends across the Mediterranean Sea and eastern Atlantic Ocean, where it is found from south west Portugal, down the African coast to Mauritania and west to Macaronesia (Alberto et al. 2008) (Fig. S1B). Congeneric species of C. nodosa are found in tropical waters of the Red Sea and Indo-Pacific, suggesting origins in the region at least prior to the closure of the Suez Isthmus, approximately 10Mya. Like C. nodosa, Cystoseira compressa has a distribution that extends across the Mediterranean and into the eastern Atlantic, where it is found west to Macaronesia and south to northwest Africa (Fig. S1C). The genus Cystoseira has recently been reclassified to include just four species with all congeneric Cystoseira spp. having warm-temperate distributions from the Mediterranean to the eastern Atlantic (Orellana et al. 2019). The distribution of Padina pavonica is conservatively considered to resemble C. nodosa and C. compressa, spanning throughout the Mediterranean and into the eastern Atlantic. We considered the poleward distribution limit of P. pavonica to be the British Isles 50ºN (Herbert et al. 2016). P. pavonica was previously thought to have a global distribution, but molecular analysis of the genus has found no evidence to support this (Silberfeld et al. 2013). Instead it has been suggested that P. pavonica was potentially misclassified outside of the Mediterranean, due to morphological similarity with congeneric species (Silberfeld et al. 2013). Padina is a monophyletic genus with a worldwide distribution from tropical to cold temperate waters (Silberfeld et al. 2013). Most species have a regional distribution, with few confirmed examples of species spanning beyond a single marine realm (sensu Spalding et al. 2007). [Metabolic rates] Net production (NP), gross primary production (GPP) and respiration (R) were measured for all species from the four sites for five different experimental temperatures containing the in-situ temperature during sampling up to a 6ºC warming (see SM Table S3 for details). Individuals of the different species were moved to methacrylate cylinders containing seawater treated with UV radiation to remove bacteria and phytoplankton, in incubation tanks at the 5 selected temperatures. Cylinders were closed using gas-tight lids that prevent gas exchange with the atmosphere, containing an optical dissolved oxygen sensor (ODOS® IKS), with a measuring range from 0-200 % saturation and accuracy at 25ºC of 1% saturation, and magnetic stirrers inserted to ensure mixing along the height of the core. Triplicates were measured for each species and location, along with controls consisting in cylinders filled with the UV-treated seawater, in order to account for any residual production or respiration derived from microorganisms (changes in oxygen in controls was subtracted from treatments). Oxygen was measured continuously and recorded every 15 minutes for 24 hours. Changes in the dissolved oxygen (DO) were assumed to result from the biological metabolic processes and represent NP. During the night, changes in DO are assumed to be driven by R, as in the absence of light, no photosynthetic production can occur. R was calculated from the rate of change in oxygen at night, from half an hour after lights went off to half an hour before light went on (NP in darkness equalled R). NP was calculated from the rate of change in DO, at 15 min intervals, accumulated over each 24 h period. Assuming that daytime R equals that during the night, GPP was estimated as the sum of NP and R. To derive daily metabolic rates, we accumulated individual estimates of GPP, NP, and R resolved at 15 min intervals over each 24 h period during experiments and reported them in mmol O2 m−3 day−1. A detailed description of calculation of metabolic rates can be found at Vaquer-Sunyer et al. (Vaquer-Sunyer et al. 2015). [Thermal distribution and thermal safety margins] We estimated the realised thermal distribution for the four experimental species by downloading occurrence records from the Global Biodiversity Information Facility (GBIF.org (11/03/2020) GBIF Occurrence Download). Occurrence records from GBIF were screened for outliers and distributions were verified from the primary literature (Alberto et al. 2008, Draisma et al. 2010, Ni-Ni-Win et al. 2010, Silberfeld et al. 2013, Telesca et al. 2015, Orellana et al. 2019) and Enrique Ballesteros (pers. comms) (Fig. S1). Mean, 1st and 99th percentiles of daily SST’s were downloaded for each occurrence site for the period between 1981-2019 using the SST products described above (Table S4). Thermal range position of species at each experimental site were standardised by their global distribution using a Range Index (RI; Sagarin & Gaines 2002). Median SST at the experimental collection sites were standardized relative to the thermal range observed across a species realized distribution, using the equation: RI = 2(SM- DM)/DB where SM = the median temperature at the experimental collection site, Dm = the thermal midpoint of the species global thermal distribution and DB = range of median temperatures (ºC) that a species experiences across its distribution. The RI scales from -1 to 1, whereby ‘-1’ represents the cool, leading edge of a species distribution, ‘0’ represents the thermal midpoint of a species distribution and ‘1’ represents the warm, trailing edge of a species distribution (Sagarin & Gaines 2002). Thermal safety margins for each population were calculated as the difference between empirically derived upper thermal limits for each population and the maximum long term habitat temperatures recorded at collection sites. Each population’s thermal safety margin was plotted against its range position to examine patterns in thermal sensitivity across a species distribution. [Growth measurements and statistical analyses] Net growth rate of seagrass shoots was measured using leaf piercing-technique (Short & Duarte 2001). At the beginning of the experiment seagrass shoots were pierced just below the ligule with a syringe needle and shoot growth rate was estimated as the elongation of leaf tissue in between the ligule and the mark position of all leaves in a shoot at the end of the experiment, divided by the experimental duration. Net growth rate of macroalgae individuals was measured as the difference in wet weight at the end of the experiment from the beginning of the experiment divided by the duration of the experiment. Moisture on macroalgae specimens was carefully removed before weighing them. Patterns of growth in response to temperature were examined for each experimental population using a gaussian function: g = ke[-0.5(TMA-μ)2/σ2], where k = amplitude, μ = mean and σ = standard deviation of the curve. Best fit values for each parameter were determined using a non-linear least squares regression using the ‘nlstools’ package (Baty et al. 2015) in R (Team 2020). 95% CI for each of the parameters were calculated using non-parametric bootstrapping of the mean centred residuals. The relationship between growth metrics and the best-fit model was determined by comparing the sum of squared deviations (SS) of the observed data from the model, to the SS of 104 randomly resampled datasets. Growth metrics were considered to display a significant relationship to the best-fit model if the observed SS was smaller than the 5th percentile of randomised SS. Upper thermal limits were defined as the optimal temperature + 2 standard deviations (95th percentile of curve) or where net growth = 0. Samples that had lost all pigment or structural integrity by the end of the experiment were considered dead and any positive growth was treated as zero. Comparative patterns in thermal performance between populations have fundamental implications for a species thermal sensitivity to warming and extreme events. Despite this, within-species variation in thermal performance is seldom measured. Here we compare thermal performance between-species variation within communities, for two species of seagrass (Posidonia oceanica and Cymodocea nodosa) and two species of seaweed (Padina pavonica and Cystoseira compressa). Experimental populations from four locations spanning approximately 75% of each species global distribution and a 6ºC gradient in summer temperatures were exposed to 10 temperature treatments (15ºC to 36ºC), reflecting median, maximum and future temperatures. Experimental thermal performance displayed the greatest variability between species, with optimal temperatures differing by over 10ºC within the same location. Within-species differences in thermal performance were also important for P. oceanica which displayed large thermal safety margins within cool and warm-edge populations and small safety margins within central populations. Our findings suggest patterns of thermal performance in Mediterranean seagrasses and seaweeds retain deep ‘pre-Mediterranean’ evolutionary legacies, suggesting marked differences in sensitivity to warming within and between benthic marine communities. [Sample collection] Sample collections were conducted at two sites, separated by approximately 1 km, within each location. Collections were conducted at the same depth (1-3 m) at each location and were spaced across the reef or meadow to try and minimise relatedness between shoots or fragments. Upon collection, fragments were placed into a mesh bag and transported back to holding tanks in cool, damp, dark conditions (following Bennett et al. 2021). Fragments were kept in aerated holding tanks in the collection sites at ambient seawater temperature and maintained under a 14:10 light-dark cycle until transport back to Mallorca, where experiments were performed. Prior to transport, P. oceanica shoots were clipped to 25 cm length (from meristem to tip), to standardise initial conditions and remove old tissue for transport. For transport back to Mallorca, fragments were packed in layers within cool-boxes. Cool-packs were wrapped in damp tea towels (rinsed in seawater) and placed between layers of samples. Samples from Catalonia, Crete and Cyprus experienced approximately 12hrs of transit time. On arrival at the destination, samples were returned to holding tanks with aerated seawater and a 14:10 light-dark cycle. [Sea temperature measurements and reconstruction] Sea surface temperature data for each collection site were based on daily SST maps with a spatial resolution of 1/4°, obtained from the National Center for Environmental Information (NCEI, https://www.ncdc.noaa.gov/oisst (Reynolds et al. 2007). These maps have been generated through the optimal interpolation of Advanced Very High Resolution Radiometer (AVHRR) data for the period 1981-2019. Underwater temperature loggers (ONSET Hobo pro v2 Data logger) were deployed at each site and recorded hourly temperatures throughout one year. In order to obtain an extended time series of temperature at each collection site, a calibration procedure was performed comparing logger data with sea surface temperature from the nearest point on SST maps. In particular, SST data were linearly fitted to logger data for the common period. Then, the calibration coefficients were applied to the whole SST time series to obtain corrected-SST data and reconstruct daily habitat temperatures from 1981-2019. [Field collections] Thermal tolerance experiments were conducted on two seagrass species (P. oceanica and Cymodocea nodosa) and two brown seaweed species (Cystoseira compressa and P. pavonica) from four locations spanning 8 degrees in latitude and 30 degrees in longitude across the Mediterranean (Fig. 1, Table S1). These four species were chosen as they are dominant foundation species and cosmopolitan across the Mediterranean Sea. Thermal performance experiments from Catalonia and Mallorca were conducted simultaneously in June 2016 for seaweeds (P. pavonica and C. compressa) and in August 2016 for seagrasses (P. oceanica and C. nodosa). Experiments for all four species were conducted in July 2017 for Crete and in September 2017 for Cyprus. Horizon 2020 Framework Programme, Award: 659246; Juan de la Cierva Formacion, Award: FJCI-2016-30728; Spanish Ministry of Economy, Industry and Competitiveness, Award: MedShift, CGL2015-71809-P; Spanish Ministry of Science, Innovation and Universities, Award: SUMAECO, RTI2018-095441-B-C21

AbstractA better understanding of how climate affects growth in tree species is essential for improved predictions of forest dynamics under climate change. Long-term climate averages (mean climate) and short-term deviations from these averages (anomalies) both influence tree growth, but the rarity of long-term data integrating climatic gradients with tree censuses has so far limited our understanding of their respective role, especially in tropical systems. Here, we combined 49 years of growth data for 509 tree species across 23 tropical rainforest plots along a climatic gradient to examine how tree growth responds to both climate means and anomalies, and how species functional traits mediate these tree growth responses to climate. We showed that short-term, anomalous increases in atmospheric evaporative demand and solar radiation consistently reduced tree growth. Drier forests and fast-growing species were more sensitive to water stress anomalies. In addition, species traits related to water use and photosynthesis partly explained differences in growth sensitivity to both long-term and short-term climate variations. Our study demonstrates that both climate means and anomalies shape tree growth in tropical forests, and that species traits can be leveraged to understand these demographic responses to climate change, offering a promising way forward to forecast tropical forest dynamics under different climate trajectories.

Global mitigation pathways play a critical role in informing climate policies and targets that are in line with international climate goals. However, it is not possible to directly compare modelled results with national inventories used to assess progress under the UNFCCC due to differences in how land-based fluxes are accounted for.National inventories consider carbon flux on managed land using an area-based approach with managed land-areas determined by nations. Emissions scenarios consider a different managed land area and are calibrated against data from detailed global carbon cycle models that account for natural (indirect) and anthropogenic (direct) fluxes separately by design. To disentangle the direct and indirect components of land-based carbon fluxes, we use a reduced complexity climate model with explicit treatment of the land-use sector, OSCAR, one of the models used by the Global Carbon Project. We find the discrepancy between model and NGHGI-based accounting methods globally to be 4.4 ± 1.0 Gt CO2 yr-1 averaged over the 2000-2020 time period, which is in line with existing estimates. We then apply OSCAR to the set of pathways assessed by the IPCC to quantify how this gap evolves over time and estimate how key mitigation benchmarks change.Across both 1.5°C and 2°C scenarios, LULUCF emissions pathways aligned with NGHGI accounting practices show a strong increase in the total land sink until around mid-century. However, the ‘NGHGI alignment gap’  decreases over this period, converging in the 2050-2060s for 1.5°C scenarios and 2070s-2080s for 2°C scenarios. The convergence is primarily a result of the simulated stabilization and then decrease of the CO2-fertilization effect as well as background climate warming reducing the overall effectiveness of the land sink, which in turn reduces the indirect removals considered by NGHGIs. These dynamics lead to land-based emissions reversing their downward trend in most NGHGI-aligned scenarios by mid-century, and result in the LULUCF sector becoming a net-source of emissions by 2100 in about 25% of both 1.5°C and 2°C scenarios.Assessing emission pathways using LULUCF definitions from national inventory accounting results in downward revisions to emissions benchmarks derived from scenarios. NGHGI-aligned pathways result in earlier net-zero CO2 emissions by around 2-5 years for both 1.5°C and 2°C scenarios, and 2030 emission reductions relative to 2020 are enhanced by about 5 percentage points for both pathway categories. When incorporating the additional land removals considered by NGHGIs, the assessed cumulative net CO2 emissions to global net-zero CO2 also decreases systematically by 15-18% for both 1.5°C and 2°C scenarios.We find that increasing removals from direct fluxes in 1.5C scenarios overtake estimated removals using NGHGI conventions in the near term. However, by midcentury, the strengthening of direct removals is balanced by weakening of indirect removals, meaning that, on average, carbon removal on land accounted for using NGHGI conventions in 1.5C scenarios results in about half of the LULUCF removals in current policy scenarios. We discuss the implications of our results for future Global Stocktakes and market mechanisms under the Paris Agreement.

Abstract Woody species' requirements and environmental sensitivity change from seedlings to adults, a process referred to as ontogenetic shift. Such shifts can be increased by climate change. To assess the changes in the difference of temperature experienced by seedlings and adults in the context of climate change, it is essential to have reliable climatic data over long periods that capture the thermal conditions experienced by the individuals throughout their life cycle. Here we used a unique cross‐European database of 2,195 pairs of resurveyed forest plots with a mean intercensus time interval of 37 years. We inferred macroclimatic temperature (free‐air conditions above tree canopies—representative of the conditions experienced by adult trees) and microclimatic temperature (representative of the juvenile stage at the forest floor, inferred from the relationship between canopy cover, distance to the coast and below‐canopy temperature) at both surveys. We then address the long‐term, large‐scale and multitaxa dynamics of the difference between the temperatures experienced by adults and juveniles of 25 temperate tree species. We found significant, but species‐specific, variations in the perceived temperature (calculated from presence/absence data) between life stages during both surveys. Additionally, the difference of the temperature experienced by the adult versus juveniles significantly increased between surveys for 8 of 25 species. We found evidence of a relationship between the difference of temperature experienced by juveniles and adults over time and one key functional trait (i.e. leaf area). Together, these results suggest that the temperatures experienced by adults versus juveniles became more decoupled over time for a subset of species, probably due to the combination of climate change and a recorded increase of canopy cover between the surveys resulting in higher rates of macroclimate than microclimate warming. Synthesis. We document warming and canopy‐cover induced changes in the difference of the temperature experienced by juveniles and adults. These findings have implications for forest management adaptation to climate change such as the promotion of tree regeneration by creating suitable species‐specific microclimatic conditions. Such adaptive management will help to mitigate the macroclimate change in the understorey layer.

AbstractAlthough the role of livestock in future food systems is debated, animal proteins are unlikely to completely disappear from our diet. Grasslands are a key source of primary productivity for livestock, and feed‐food competition is often limited on such land. Previous research on the potential for sustainable grazing has focused on restricted geographical areas or does not consider inter‐annual changes in grazing opportunities. Here, we developed a robust method to estimate trends and interannual variability (IV) in global livestock carrying capacity (number of grazing animals a piece of land can support) over 2001–2015, as well as relative stocking density (the reported livestock distribution relative to the estimated carrying capacity [CC]) in 2010. We first estimated the aboveground biomass that is available for grazers on global grasslands based on the MODIS Net Primary Production product. This was then used to calculate livestock carrying capacities using slopes, forest cover, and animal forage requirements as restrictions. We found that globally, CC decreased on 27% of total grasslands area, mostly in Europe and southeastern Brazil, while it increased on 15% of grasslands, particularly in Sudano‐Sahel and some parts of South America. In 2010, livestock forage requirements exceeded forage availability in northwestern Europe, and southern and eastern Asia. Although our findings imply some opportunities to increase grazing pressures in cold regions, Central Africa, and Australia, the high IV or low biomass supply might prevent considerable increases in stocking densities. The approach and derived open access data sets can feed into global food system modelling, support conservation efforts to reduce land degradation associated with overgrazing, and help identify undergrazed areas for targeted sustainable intensification efforts or rewilding purposes.

Abstract Background The global human footprint has fundamentally altered wildfire regimes, creating serious consequences for human health, biodiversity, and climate. However, it remains difficult to project how long-term interactions among land use, management, and climate change will affect fire behavior, representing a key knowledge gap for sustainable management. We used expert assessment to combine opinions about past and future fire regimes from 99 wildfire researchers. We asked for quantitative and qualitative assessments of the frequency, type, and implications of fire regime change from the beginning of the Holocene through the year 2300. Results Respondents indicated some direct human influence on wildfire since at least ~ 12,000 years BP, though natural climate variability remained the dominant driver of fire regime change until around 5,000 years BP, for most study regions. Responses suggested a ten-fold increase in the frequency of fire regime change during the last 250 years compared with the rest of the Holocene, corresponding first with the intensification and extensification of land use and later with anthropogenic climate change. Looking to the future, fire regimes were predicted to intensify, with increases in frequency, severity, and size in all biomes except grassland ecosystems. Fire regimes showed different climate sensitivities across biomes, but the likelihood of fire regime change increased with higher warming scenarios for all biomes. Biodiversity, carbon storage, and other ecosystem services were predicted to decrease for most biomes under higher emission scenarios. We present recommendations for adaptation and mitigation under emerging fire regimes, while recognizing that management options are constrained under higher emission scenarios. Conclusion The influence of humans on wildfire regimes has increased over the last two centuries. The perspective gained from past fires should be considered in land and fire management strategies, but novel fire behavior is likely given the unprecedented human disruption of plant communities, climate, and other factors. Future fire regimes are likely to degrade key ecosystem services, unless climate change is aggressively mitigated. Expert assessment complements empirical data and modeling, providing a broader perspective of fire science to inform decision making and future research priorities.